Phylogeny and biogeography of the Platystictidae (Odonata) Tol, J. van

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Note: To cite this publication please use the final published version (if applicable). 5. Revision of the Platystictidae of the Philippines (Odonata), excluding the Drepanosticta halterata-group, with descriptions of twenty-one new species

J. van Tol

Tol, J. van. Revision of the Platystictidae of the Philippines the Moluccas and New Guinea. Several species confined to (Odonata), excluding the Drepanosticta halterata group, with Palawan have sister-group relationships with species from descriptions of twenty-one new species. Borneo. The affinities of various other species confined to Zool. Meded. Leiden 79-2 (10), 22.vii.2005: 195-282, figs the Sulu archipelago, are unsettled as yet. The species of 1-109, 1 table. – ISSN 0024-0672. Platystictidae here assigned to Protosticta Selys are presumably not closely related to the type species, P. simplicinervis Key words: Odonata; Platystictidae; Drepanosticta; Protosticta; Selys from Sulawesi. However, a better placement has to Sulcosticta; Philippines; new species; new genus. await a more detailed phylogenetic study of the family. For Thirty-one species of the family Platystictidae of the three species the new genus Sulcosticta gen. n. is erected. Philippines are revised, i.e. all species recognised, excluding These species are closely allied based on the structure of the the species of the Drepanosticta halterata-group. The following appendages, but should have been assigned to different genera new taxa are described: 16 species in Drepanosticta Laidlaw: if based on the present generic definitions. D. acuta sp.n., D. aurita sp. n., D. centrosaurus sp. n., D. Many species here described have small distributional ranges, clados sp. n., D. flavomaculata sp. n., D. furcata sp. n., D. a common phenomenon in Platystictidae. Since most forests hermes sp. n., D. krios sp. n., D. luzonica sp. n., D. malleus sp. in the Philippines are heavily under threat or have already n., D. myzouris sp. n., D. paruatia sp. n., D. pistor sp. n., D. disappeared in the last fifty years, several taxa described in quadricornu sp. n., D. rhamphis sp. n., D. trachelocele sp. n., this paper should be considered under threat of immediate two in Protosticta Selys, viz. P. lepteca sp. n. and P. plicata sp.n., extinction. and three in Sulcosticta gen. n., viz. S. striata sp. n., S. pallida sp. n. and S. viticula sp. n. The status of eleven previously discribed nominal taxa is established. One, D. septima Introduction Needham & Gyger, is doubtfully considered a synonym of D. Forest damselflies, Platystictidae, are among the most mylitta Cowley. characteristic elements of seepage areas, trickles and Based on a preliminary phylogenetic analysis, the species of Drepanosticta are divided into informal species groups. Most small streams of virgin forests in Southeast Asia. species of the Philippines have affinities to species of Sulawesi, Both the mainland and many of the islands of this region are inhabited by great variety of species of the genera Protosticta Selys and Drepanosticta Laidlaw Previously published in: Zoölogische Mededelingen 79-2 (Fig. 1). Most platystictid species of Southeast Asia [2005]: 195-282, figs. 1-109, table 1. were described by Lieftinck (e.g. 1937, 1939, 1961,

131 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

Figure 1. Drepanosticta halterata complex. Habitus of male. The species of theD. halterata species group are not fully treated in this paper.

1965), while more recently mainly Asahina (e.g. 1984), species in Drepanosticta and three species in Protosticta Hämäläinen (1991, 1999), Wilson (1997), Wilson awaited description. & Reels (2001) and Van Tol (2000) have contributed The paper by Hämäläinen & Müller (1997) was to the regional knowledge of this family. For the based on a careful study of the specimens available Philippines, especially the publications by Needham in museums, but focussed on the collection brought & Gyger (1939, 1941) are relevant, although the first together by Mr. Roland A. Müller (St. Gallen, Philippine species were already described by Brauer Switzerland) and his collaborators between 1985 (1868). and 1997 (Fig. 2). The Müller collection is the first Drepanosticta and Protosticta species generally have collection of Philippine Odonata representative of small distributional ranges, and island endemicity at all parts of the country. It is also outstanding in the the species level is high, e.g. on Sulawesi for the genus documentation of localities. The National Museum of Protosticta it is hundred percent (Van Tol, 2000). Natural History Naturalis at Leiden and Mr. Roland The same picture of small ranges and high island A. Müller were able to reach an agreement on the endemicity is apparent for platystictids of many other deposition of this collection in Leiden in 1998. The islands of Indonesia, and is also true for species of this collection is rich in species of previously neglected damselfly family of the Philippines as well. Hämäläinen families as Platycnemididae and Platystictidae. Some & Müller (1997) mentioned the high number of presently still undescribed species are represented species, especially in the genus Drepanosticta. They with more than 200 specimens. Nearly all material enumerated 35 species in the family, 32 species of the Müller collection was identified by Dr Matti of Drepanosticta and three species of Protosticta. Hämäläinen (Espoo, Finland). His pioneering work Only twelve valid Philippine species in the genus has revealed the value of this collection in many ways. Drepanosticta have been described so far, and one other Hämäläinen himself published a series of papers nominal taxon is considered a synonym. So, twenty describing taxa new to science (see Hämäläinen

132 Chapter 5 Platystictidae of the Philippines

& Müller, 1997 for an overview; and Hämäläinen 1997, 2000; Gassmann & Hämäläinen 2002, Van Tol & Müller, 2003 for results published since that time). This paper is part of a series of papers in which I hope to revise the Platystictidae of the Oriental and Papuan regions, and to reconstruct the phylogeny of this family, including the Palaemnema species of the New World. New species of Drepanosticta from various parts of the mainland of Southeast Asia, and from several islands in Indonesia, await description. Also, the generic division of the Oriental species is far from satisfactory, as it is mainly based on one wing venational character, which seems to contradict a grouping of species based on an analysis including various other characters (see also Orr, 2003: 69-72). The present papers includes all Philippine species of Platystictidae, except the species of the Drepanosticta halterata group. The latter group is widespread in Luzon (incl. Batan), the West Visayas (Panay, Negros, Cebu, Sibuyan) and East Visayas (Samar, Bohol). Although material is available in sufficient numbers, and clearly recognisable variation in characters exists between populations, the Figure 2. Approximate position of localities of the specimens in the distinction of species needs further study of Roland Müller collection (from: Hämäläinen & Müller 1997: 250). Scale the material, including some type specimens. bar in kilometers. Three obviously closely related species are assigned here to a new genus, as a first step to define monophyletic groups within this family. reconstruct the phylogenetic relationships have only The studies of the forest damselflies of Southeast Asia just started. by others and myself, partly still unpublished, slowly The uneasy observation must be made that many of the unravel a fascinating image of diversity as a result of spectacular life forms described in this paper are on the millions of years of evolution. Most species are dull- verge of extinction. Several species are described here coloured, small and inconspicuous insects, superficially from only one or very few specimens collected from looking very similar. The variation in small details of a forest reserve of one hectare of less. Since several of the male anal appendages is remarkable in the species such precious sites already got lost since the collections of the mainland. Most species of the Philippines have were made, we must fear for their survival. One may structurally similar anal appendages in the male, but only be hopeful that the present paper may contribute they show the most exuberant structures in projections to the awareness in the Philippines of the huge, unique of the pronotum. The information of these features to and irreplaceable diversity of insect life of this country.

133 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

Material and methods are presented separately. Illustrations are based on specimens in the RMNH collection (except for the This paper is based on the material in the National holotype of Drepanosticta megametta Cowley); the Museum of Natural History (Leiden), including the references to the actual specimens are given in the former collections of Roland A. Müller (St. Gallen, captions. The distributions are summarized at the end Switzerland) (see above) and Dr. Matti Hämäläinen of each description. A series of distribution maps is (Espoo, Finland). The latter collection, although presented at the end of the paper (Figs 99-109). much smaller than the Müller collection, was also valuable since it includes both duplicates from the Phylogenetic relationships Müller collection retained after his identifications, as well as specimens collected by Hämäläinen Although a more elaborate analysis of the phylogeny of himself. Types were examined, if necessary, for several the Philippine Platystictidae has to await the analysis previously described species. The study of the types of all species of this family for Southeast Asia, a few of species described by Needham and Gyger had to remarks can be made. be postponed. I have relied on the descriptions, or Firstly, various species of Palawan and Busuanga material in the Leiden museum identified by M. A. have little in common with most other species of the Lieftinck after comparison with types. Philippines. Species such as D. ceratophora Lieftinck Deposition of other material is documented, with and D. paruatia sp. n. have sister-group affinities with collections identified by codens as follows: BMNH, species of Borneo. The relationship of another species Natural History Museum, London, UK [= British of the Palawan area, D. quadricornu sp. n., is uncertain. Museum (Natural History)]; CASC, California Based on the remarkably similar structure of the Academy of Sciences, San Francisco, USA; CUIC, male anal appendages of almost all other species of Cornell University, Ithaca, New York, USA; FMNH, Philippine Drepanosticta, these species are presumably Field Museum Natural History, Chicago, Illinois, derived from one common ancestor. Based on the USA; IRSN, Institut royal des Sciences Naturelles de structure of both anal appendages and pronotum, the Belgique, Bruxelles, Belgium; MCZC, Museum of closest affinities of this group are definitely with species Comparative Zoology, Cambridge, Massachusetts, as D. ephippiata Lieftinck from Sulawesi, or D. clavata USA; MNHN, Museum National d’Histoire Lieftinck from New Guinea (see also Lieftinck 1937: Naturelle, Paris, France; RMNH, National Museum 72-74). The males of the Philippine species, although of Natural History, Leiden, The Netherlands [formerly also showing subtle differences in the structure of Rijksmuseum van Natuurlijke Historie]; SMFD, the anal appendages, are most easily distinguished by Senckenberg Museum, Frankfurt a. M.; ZMUC, the structure of the pronotum. Both the anterior and Zoological Museum University of Copenhagen. the posterior lobe of the pronotum can be furnished Descriptions of all species start with a diagnosis, with the most exuberant projections, sometimes even enumerating the most distinct characters as compared exceeding in length the size of the pronotum itself. The to other Philippine species, especially in the species characters of the pronotum as mentioned here are those groups distinguished. Terminology generally follows of the male. In some species male and female have very Watson & O’Farrell (1991), and Cowley (1936) for similar projections on the pronotum, while seemingly specialised terminology of the head structure. For most closely related species male and female are structurally species the description of the male is based on the different in this respect. holotype. Measurements are also for the holotype, with Within this group of closely related species, various variation of the paratypes in brackets. The descriptions species groups could be identified based on a of the females are concise, typically comparing the preliminary analysis of the characters. These groups female with the male. Measurements for the females have been used in this publication to provide a first

134 Chapter 5 Platystictidae of the Philippines

structuring of the variation. Some more widespread the group of ‘geographically recognizable forms’ of species show clinal variation in the structure of e.g. the D. mylitta Cowley was not split, but kept under the projections of the pronotum. Also, some islands forms name D. mylitta. Further studies, including molecular were recognisable. Decisions whether such forms had studies, are needed to provide an understanding of the to be considered as separate species, were difficult and relationships of these complexes, thus also providing a are debatable. For instance, the ‘form’ of D. belyshevi better basis for such taxonomic decisions. Hämäläinen from Samar was considered so distinct Various species in this paper are assigned to the genus that it is raised here to species level. On the other hand, Protosticta Selys, based on the absence of the anal bridge vein (Ab) at the base of the wing. However, it is unlikely that these species are closely Figure 3. Faunal regions in the Philippines, with names of islands related to the type species of Protosticta, P. mentioned in the text. Abbreviations for regions as follows: L: Luzon simplicinervis Selys from Sulawesi. It is certain region, MDO: Mindoro region, WV: West-Visayan region, EV: East that the venational character distinguishing Visayan subregion, MNO: Mindanao subregion (EV plus MNO together Protosticta and Drepanosticta will prove to form Mindanao region), P: Palawan group (part of Greater Sunda region), S: Sulu rgion. Luzon, Mindoro, West-Visayan and Mindanao regions be of little phylogenetic significance. It has comprise the Philippine biogeographic region proper (from: Hämäläinen to remain open for further research whether & Müller 1997: 251). Scale bar in kilometers. all Philippine ‘Protosticta’ belong to the Drepanosticta-complex of the Philippines, or should be regarded as a distinct genus. Unfortunately, all species of this group of species are rare in the collection, and presently no material is available for molecular analysis. Such an analysis may also provide further insight in the relationships of the species here assigned to the new genus Sulcosticta. The generic placement of most species in the genus Drepanosticta in the present sense seems rather unproblematic. However, the genus is based on a synplesiomorphy, and its monophyly is both unestablished as well as unlikely. A further division into monophyletic groups has to await the revision of the species of the mainland of South-East Asia. The assignment of other species to Protosticta is, as explained above, preliminary. The character presently defining the genus Protosticta, viz. the absence of the Ab vein, is clearly not a unique apomorphy in the family, and will prove unsuitable for the definition of a monophyletic group. For example, one of the species here included in Protosticta on the characters of the male would have to be included in Drepanosticta if

135 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

Table 1 Distribution of Platystictidae described in this paper, per faunal region. The East Visayan and Mindanao regions are usually considered subregions of one single region. × Present. — o Absent. — (×) Status uncertain. — ? Locality uncertain. Luzon Mindoro Visayan West East Visayan Mindanao Sulu Palawan Drepanosticta acuta sp. n. ×oooooo Drepanosticta aries Needham & Gyger oooo×oo D. krios sp. n. o o o ? × × o D. rhamphis sp. n. ×oooooo D. belyshevi Hämäläinen o o o × o o o D. flavomaculata sp. n. o o o o × o o D. trachelocele sp. n. o o o × o o o D. luzonica sp. n. ×oooooo D. moorei Van Tol & Müller × oooooo D. clados sp. n. o o o o × o o D. furcata sp. n. oo×oooo D. hermes sp. n. o o o o × o o D. lymetta Cowley oooo×oo D. taurus Needham & Gyger oooo×oo D. centrosaurus sp. n. oooo×oo D. megametta Cowley oooo×oo D. aurita sp. n. o×ooooo D. ceratophora Lieftinck oooooo× D. malleus sp. n. o o o o × o o D. mylitta Cowley (×) o o × (×) o o D. myzouris sp. n. ×oooooo D. paruatia sp. n. oooooo× D. pistor sp. n. ×o×oooo D. quadricornu sp. n. oooooo× Protosticta annulata Selys × oooooo P. lepteca sp. n. ×oooooo P. plicata sp. n. oo×oooo Sulcosticta pallida sp. n. ×oooooo S. striata sp. n. ×oooooo

Total 11 1 3 3-4 10-11 1 3

based on the wing venation of the female. genitalia. It is expected that a further phylogenetic Three species with different wing venation, but analysis may reveal that more species will have to be otherwise evidently closely related, are placed in a placed in this genus. new genus based on the similar structure of the male

136 Chapter 5 Platystictidae of the Philippines

Distributional patterns peninsula of Mindanao had a more western position at the northern part of the Celebes Sea. In the This is not the place to provide a profound late Miocene, the island of Luzon arc still moved zoogeographical analysis based on the Platystictidae, northward, and partly collided with northern Palawan. since the prerequisites for such an analysis are not Also the during the Pliocene the clockwise rotation of fulfilled. Nevertheless, some brief remarks on the the plate continued. As a result, for instance, the island distributions of presumed monophyletic groups, or of Halmahera was still north of the Vogelkop peninsula sister-group relationships with other areas in South- in the Pliocene, 500 km east of its present position. East Asia, may contribute to the definition of a more The island of Luzon, in its present shape, was only detailed zoogeographical study. The platystictids will be formed during the late Pliocene (2 Ma). discussed based their occurrence in the faunal regions Another important, and geologically recent, as defined by e.g. Vane-Wright (1990) (see Fig. 3). phenomenon is the lowering of the sea level during The biogeography of the Philippines has been discussed the ice ages of the Quaternary. Many of the island several times the last decades, also in relation to the were connected when sea level was c. 150 m lower palaeogeography of the region. A relevant discussion is than today. The channel between Borneo and the De Jong & Treadaway (1993: 81-111). Their study was Philippines is not deeper than 145 m and Borneo and hampered by insufficient knowledge of the phylogeny Palawan formed one island, while also many of the of the Hesperiidae, making a cladistic biogeographical Sulu islands were connected at the time. analysis impossible as well. Instead, they compared As discussed above, the Palawan group of species seems hypothesized distribution patterns derived from to be most closely related to the species of Borneo, geological events with actual patterns found. If such and not to the other species of the Philippines. This patterns cannot be traced, dispersal rather than a phenomenon is well known in other groups as well, geological event was supposed to be the cause of such and can be understood with present knowledge of a pattern. Mainly based on terrestrial vertebrates, recent geology. It should, however, be realised that the Philippines were subdivided in faunal regions. various lineages with many autapomorphies living in Interestingly, this subdivision is similar to the regions as Palawan have not reached Borneo, or have become defined by Semper (1892) based on butterflies around extinct in Borneo. hundred years earlier. The area of highest diversity in the Platystictidae (table 1) is the eastern part of the island of Mindanao. Many The palaeogeography of the region is very complicated species widespread in Mindanao, and in some cases indeed. Continental fragments are restricted to the also in other islands, occur in this area together with western part of the Philippines, while most islands many species with extremely small distributional ranges were formed along the Philippine plate during the and endemic to this area. This is best shown in the D. last 40 million years. These islands were formed along lymetta group. the westside of the Philippine Sea Plate by closewise The fauna of Luzon is most diverse on the level of rotation of the plate from the late Eocene (42 Ma). species-groups. The belyshevi group, the halterata Further to the west, not far from the mainland, parts group, the moorei group, the Protosticta group and of Palawan, Panay and Mindoro (Calamian block) several unplaced species are all confined to, or strongly were situated. The Calamian block rifted eastward by represented in, Luzon. From most other islands only seafloor spreading during the Oligocene. Even during one or very few species are known. Although this the Miocene, the islands arc along the westside of the observation may partly be an artifact due to insufficient Philippine Sea Plate still existed, including (parts of) collecting, or massive deforestation in some parts of the Luzon, Zambales, East Mindanao, North Sulawesi Philippines in recent times, it is in agreement with the and (more to the east) Halmahera. The Zamboanga generally recognized biogeographical pattern. The small

137 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

number of species on Mindoro, however, is remarkable Laidlaw) ...... 6 and should be studied further. – Anal bridge absent (Protosticta Selys) ...... 31 6 (5) Posterior margin of posterior lobe of pronotum Systematic part with a single, median process, curved anteriad (Fig. 55) ...... D. ceratophora Lieftinck Key to the males of Platystictidae of the Philippine – Posterior margin of posterior lobe of pronotum Islands simple without processes, or with a pair of shorter or longer processes ...... 7 1. Inferior appendages (ventral view) broad and short, 7. Anterior and posterior lobes with conspicuous, the top bifid or with top shining brown sclerotized paired processes on hind margin, the processes and flattened (Fig. 95); synthorax with variegated distinctly longer than length of lobes themselves pattern of longitudinal stripes (Sulcosticta gen. n.) ...... 8 ...... 2 – Anterior or posterior lobes (not both) with – Inferior appendages much broader at base than at conspicuous processes longer than length of lobes top, usually base stout, the distal half much more themselves, or processes lacking on both anterior slender, without shining brown sclerotization at and posterior lobes ...... 11 distal margin; synthorax usually uniform brown 8. Transverse occipital carina poorly developed, at least or with one longitudinal stripe; if synthorax with without conspicuous angulate lateral extremities 9 variegated pattern , then inferiors long and slender – Transverse occipital carina well developed with ...... 5 distinct, angulate lateral extremities ...... 10 2. Inferior appendage undivided, the top flattened and 9 (8). Paired processes of anterior lobe of pronotum sclerotized, directed dorsad, the margin with a row filiform, much more slender than the processes of teeth (Polillo) .....Sulcosticta sp. n. [unnamed] of posterior lobe (Fig. 28); hind wing c. 22 mm – Inferior appendage bifid, or with distinct inward (Luzon) ...... D. moorei Van Tol & Müller directed projection ...... 3 – Paired processes of anterior lobe of pronotum 3. Inferior appendage subterminally with distinct, flat and triangular; paired processes of posterior inward directed projection ending as flat sclerotized lobe hornlike (Figs 77-78); hind wing c. 18 mm surface ...... S. viticula sp. n. (Palawan, Busuanga) ...... D. quadricornu sp. n. – Inferior appendage bifid, both parts subequal, 10 (8) Base of posterior processes of posterior lobe of inner part sclerotized, but not ending in a flat pronotum (dorsal view) very close together, distance surface ...... 4 between both processes c. two times the width of 4. Synthorax very pale, mesepisternum greyish process; processes smoothly curved abaxiad (Fig. 16) yellow, except along dorsal carina; anterior lobe of ...... D. belyshevi Hämäläinen prothorax simple without processes; Anal bridge – Base of posterior processes wide apart, distance present but not connected with Anal crossing .... between both processes more than five times the ...... S. pallida sp. n. width of process; processes curved somewhat – Synthorax with a complex pattern of dark and pale dorsad, the distal three-quarters ventrad (Fig. 22) . markings, mesepisternum predominantly black with ...... D. trachelocele sp. n. a narrow pale stripe against humeral suture; anterior 11 (7) Transverse occipital carina poorly developed, lobe of prothorax with a paired of long processes, at least without conspicuous angulate lateral curved anteriad, twice as long as length of anterior extremities; hind wing 25 mm or more ...... 12 lobe itself ...... S. striata sp. n. – Transverse occipital carina with distinct lateral 5 (1). Anal bridge (Ab) present and connected extremities; hind wing shorter than 25 mm ....15 with Anal crossing as Y-shaped vein (Drepanosticta 12 (11) Hind margin of posterior lobe of pronotum

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with a pair of processes, longer than the median 17. Synthorax castaneous; lateral corners of posterior length of the posterior lobe ...... 13 lobe of pronotum sharply projected (Fig. 64). . . . . – Hind margin of posterior lobe of pronotum with ...... D. mylitta Cowley smoothly rounded, or somewhat lifted, but without – Synthorax brownish black; lateral corners of a process longer than median line of posterior lobe posterior lobe of pronotum rounded, or distinct ...... 14 processes longer than median length of posterior 13. Tip of inferior appendage with a long subterminal, lobe itself on hind margin ...... 18 medially directed tooth (Figs 69-70); processes of 18. Posterior lobe of pronotum with lateral corners posterior lobe with top widening as a funnel, top broadly rounded (‘ears’), the projections densely set flat as a sucker (Fig. 68) ...... D. myzouris sp. n. with long setae (Fig. 52) ...... D. aurita sp. n. – Tip of inferior appendage on innerside smooth or – Posterior lobe of pronotum with distinct processes, somewhat projected, but without a distinct tooth; longer than the median line of the lobe ...... 19 processes of posterior lobe tapering or club-shaped, 19. Posterior lobe of pronotum with paired collar-like but tip not flat ...... D. halterata group processes, broadly rounded at top, constricted at

base and broadly connected with the lobe .....20 The following described species are included in this group: D. halterata Brauer, D. philippa Lieftinck, and – Posterior lobe of pronotum with paired processes, D. trimaculata Lieftinck; several undescribed species are usually slender, the top in some species split into available in collections. The status of various nominal branches or hammer-like structure ...... 21 taxa in uncertain. This group is common and widespread 20. Paired processes of posterior lobe of pronotum in Luzon. A special publication on this group is in approximately as long as wide, only somewhat preparation. constricted in the middle (Figs 49-50) ...... 14 (12) Superior appendages with distinct dorsal tooth ...... D. megametta Cowley (Figs 59-60); lateral sides of hind lobe of pronotum – Paired processes of posterior lobe of pronotum flat and sharp (Fig. 58); head very smooth and much wider than long, the middle distinctly shining, without microsculpture (c. 30 times constricted, thus the top with long projections (Fig. magnification) ...... D. lestoides Brauer 46) ...... D. centrosaurus sp. n. – Superior appendages without distinct ventral tooth 21 (19). Paired processes of posterior lobe of pronotum (Figs 26-27); lateral sides of hind lobe of pronotum curved, undivided, usually approximately as long as somewhat uplifted and broadly rounded (Fig. 25); the length of posterior lobe itself (D. aries group) . head coriaceous ...... D. luzonica sp. n...... 22 15 (11) Metepisternum and metepimeron – Paired processes of posterior lobe of pronotum predominantly creamish white; mesepisternum with erect, not distinctly curved, but tip frequently split a bluish antehumeral stripe ...... and developed as a knob, or a fork, or a disk ...24 ...... Drepanosticta paruatia sp. n. 22. Posterior part of metepisternum and metepimeron – Metepisternum and metepimeron brown; pale with creamish yellow squarish markings; abdominal coloration, if present, a squarish, creamish white segment 10 and appendages pale (Figs 20-21) .... marking against posterior margin of synthorax; no ...... D. flavomaculata sp. n. antehumeral stripe ...... 16 – Posterior part of metepisternum and metepimeron 16. Synthorax with a complex coloration of brownish dark brown, concolorous with other parts of black and pale markings; superior appendages in synthorax; at least segment 10 brown, colour not lateral view strongly curved (Figs 75-76) ...... not different from segment 9 ...... 23 ...... D. pistor sp. n. 23. Px 16-17 in fore wing, 15-16 in hind wing – Synthorax concolorous; superior appendages in (Mindanao, Mt Apo) . D. aries Needham & Gyger lateral view nearly straight ...... 17 – Px 14-15 in fore wing, 13-14 in hind wing

139 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

(Mindanao, but not in Mt. Apo region) ...... – Synthorax brownish black; paired processes of ...... D. krios sp. n. posterior lobe of pronotum divided into two longer, 24 (21). Superior appendages in lateral view curved, usually asymmetrical branches which are more the inferiors distinctly surpassing the level of the slender than the stem (Mindanao Id) ...... 30 superiors (Figs 32-33) ...... D. clados sp. n. 30. Distal half of inferior appendage curved like as – Superior appendages in lateral view more or less clasper; outer branch of paired processes of posterior straight, the inferiors and superiors in lateral view lobe of pronotum distinctly longer than inner approximately of equal length ...... 25 branch (Fig. 37) ...... D. hermes sp. n. 25. Top of paired processes of posterior lobe of – Distal half of inferior appendage straight; inner pronotum ending abruptly in a smaller or larger branch of paired processes of posterior lobe of hook ...... 26 pronotum typically longer than outer branch, but in – Top of paired processes straight or split, but some specimens difference indistinct (Fig. 40) ... branches of splitting not forming an angulate hook ...... D. lymetta Cowley with main stem of process ...... 28 31 (5). Anterior lobe of pronotum with paired 26. Paired processes of posterior lobe of pronotum processes, approximately as long as median line of very long and heavy, length more than twice the anterior lobe itself; hind lobe of pronotum with a median line of the posterior lobe itself; branches of pair of longer processes, which are curved anteriad T-splitting also strongly built (Fig. 61); Y-vein at (Fig. 84) ...... Protosticta lepteca sp. n. base of wing distinctly stalked ....D. malleus sp. n. – Anterior lobe of pronotum simple, without – Paired processes of posterior lobe of pronotum processes; hind lobe with short processes never bent approximately as long as median line of posterior anteriad ...... 32 lobe itself; Y-vein (sub)-sessile ...... 27 32. Paired processes of hind lobe of pronotum erect, 27. Superior appendage (dorsal view) with a very short, the top sharp (Fig. 81) ...... P. annulata (Selys) mediad tooth near appendage base, and a hardly – Paired processes of hind lobe of pronotum with discernable tubercle at base of club-shaped part of rounded top, formed since the process is halfway appendage; paired posterior processes of posterior recurved distad the fully 180˚ (Fig. 87) ...... lobe of pronotum with distinct stem (Fig. 13) ...... P. plicata sp. n. (Catanduanes Id.) ...... D. rhamphis sp. n. – Superior appendage with a long and acute, ventro- List of Platystictidae of the Philippines mediad tooth at base of club-shaped top; paired posterior processes of posterior lobe of pronotum The species are arranged by species-group. The club-shaped part with short stem (Fig. 4) (Luzon) sequences of the species-groups, and the species within ...... D. acuta sp. n. the species-groups, are alphabetical. 28 (25). Paired processes of posterior lobe of pronotum straight and erect (perpendicular to body axis), the Drepanosticta Laidlaw, 1917 top somewhat thicker (Fig. 43) ...... D. aries group ...... D. taurus Needham & Gyger 1. D. acuta sp. n...... 141 – Paired processes of posterior lobe of pronotum split 2. D. aries Needham & Gyger, 1941 ...... 143 in branches, usual of unequal length, the process 3. D. krios sp. n...... 144 not 90˚ erect, but more directed posteriad ....29 4. D. rhamphis sp. n...... 146 29. Synthorax castaneous; paired processes of posterior D. belyshevi group lobe of pronotum divided in two short branches 5. D. belyshevi Hämäläinen, 1991 ...... 147 similar to the stem (Fig. 34) (Siquijor Id) ...... 6. D. flavomaculata sp. n...... 149 ...... D. furcata sp. n. 7. D. trachelocele sp. n...... 151

140 Chapter 5 Platystictidae of the Philippines

[D. halterata group] straight, rather than fractured, IR3 vein, and the [not reviewed in this paper] presence of an Anal bridge, joining the Anal crossing or D. halterata Brauer, 1868 the hinder margin of the wings. Here, a new genus is D. philippa Lieftinck, 1961 erected for three Philippine species based on the shape D. trimaculata Lieftinck, 1939 of the superior appendages. One of these species does D. moorei group have an Anal bridge, which does not join the Ac before 8. D. luzonica sp. n...... 152 or at the wing margin (‘sessile Y-vein’), but meets the 9. D. moorei Van Tol & Müller, 2003 ...... 154 hinder margin of the wing far from the Ab. D. lymetta group The Philippine species of Drepanosticta are placed 10. D. clados sp. n...... 155 here in various informal groups based on preliminary 11. D. furcata sp. n...... 157 phylogenetic studies, as explained above. The following 12. D. hermes sp. n...... 158 groups have been defined (in alphabetical order): D. 13. D. lymetta Cowley, 1936 ...... 159 aries group, D. belyshevi group, D. moorei group, D. 14. D. taurus Needham & Gyger, 1941 .... 161 lymetta group, D. megametta group. The groups are D. megametta group named after the species of the group that was first 15. D. centrosaurus sp. n...... 163 described. 16. D. megametta Cowley, 1936 ...... 164 Drepanosticta aries group Drepanosticta: other species 17. D. aurita sp. n...... 165 Transverse occipital carina well developed, with distinct 18. D. ceratophora Lieftinck, 1974 ...... 167 lateral extremities; anterior lobe of pronotum smooth 19. D. lestoides Brauer, 1868 ...... 168 and flat; posterior lobe of pronotum with flat and 20. D. malleus sp. n...... 170 curved processes, approximately the median length of 21. D. mylitta Cowley, 1936 ...... 172 the posterior lobe; synthorax brown; pterostigma wider 22. D. myzouris sp. n...... 174 than high; superior appendages with dorsal tooth; 23. D. paruatia sp. n...... 175 inferior appendages straight, the tip curved medio- 24. D. pistor sp. n...... 176 dorsad. 25. D. quadricornu sp. n...... 178 Included species: D. acuta sp. n., D. aries Needham & Gyger, D. krios sp. n., D. rhamphis sp. n. Protosticta Selys, 1885 Distribution: Luzon, Catanduanes, Mindanao, Sulu 26. P. annulata Selys, 1886 ...... 180 archipelago; Samar doubtful. 27. P. lepteca sp. n...... 182 28. P. plicata sp. n...... 183 Drepanosticta acuta van Tol, sp. n. (Figs 4-6, 99) Sulcosticta gen. n. 29. S. pallida sp. n...... 185 Drepanosticta sp. n. 10. – Hämäläinen & Müller 1997: 257, 30. S. striata sp. n...... 185 277 (Luzon). 31. S. viticula sp. n...... 188 Type material . – Holotype male [JvT 18985] in RMNH: Unplaced specimens ...... 189 ‘Philippines. Luzon Id. Camarines Sur. Pili. Bungcao Curry. Mt. Isarog. Caririca river. 200-400 m. 4-15 Aug 1997, Celso Nazareno’. – Paratypes (all Philippine Islands). Luzon: Mt. Drepanosticta Laidlaw Isarog, 8.iv.1916 (G. Boettcher) 1 female (in SMFD); P. I., Drepanosticta Laidlaw, 1917: 339. M.T. Iriga / Camarines Sur, 6.iv.1962 (H.M. Torrevillos) 1 male; Camarines Sur. Pili. Bungcao Curry. Mt. Isarog. Drepanosticta Laidlaw is traditionally distinguished Caririca river. 200-400 m, 4-15.viii.1997 (Celso Nazareno) 8 within the Platystictidae by the combination of a males 4 females.

141 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

4 5 6

Figures 4-6. Drepanosticta acuta sp. n., male [JvT 18981, Luzon, Camarines Sur, Pili, Bungcao Curry, Mt Isarog, 200-400 m, 4-15 Aug 1997]. – 4, pronotum, oblique view. – 5, anal appendages, dorsal view. – 6, idem, left lateral view. Scale bar 1 mm.

Diagnosis. – Medium-sized; very broad superior top directed 90˚ abaxiad, as head of woodpecker, distal appendages in lateral view, which also have a part with long setae. Synthorax unicolorous bronze- remarkably long and sharp ventro-mediad tooth at the black, especially metepisternum and metepimeron base of the widened part. The three other species in this shining black, rest coriaceous. Legs dirty yellow, femora group with short to very short, inconspicuous tooth near joints of tibiae with brownish black ring. Wings at base of widened part. Females without projections hyaline; venation brown, somewhat darker in posterior on posterior margin of posterior lobe of pronotum; part; Px 17 in fore wing, Px 16 in hind wing; R4+5 the same structure in D. rhamphis sp. n. with distinct arising just distal to subnodus, IR3 just distal to that projections. level; Arculus arising just distal to Ax2; quadrangle somewhat widening posteriorly in fore wing; Male [holotype, JvT 18985]. – Head. Labrum, pterostigma castaneous, c. two times as wide as high, mandibles and anteclypeus bluish white; anterior proximal side acutely angulate; cells between Costa and border of labrum with narrow black line; rest of head R1 distal to pterostigma undivided. bronze-black, postclypeus and frons shining brownish Abdomen. Segments 1-7 brown to brownish black, black, dorsal side of head coriaceous, with many with distinct pale yellow basal annulae covering 1/8 to elongate striae lateral to ocelli; transverse occipital 1/5 of length of each segment; segments 8-10 brownish carina with distinct acutely angulate extremities; black. Appendages (Figs 5-6) with superiors stout at antenna with scapus brownish black, pedicellus base, distal three-fifths tapering, a distinct ventro- yellowish white, tip of pedicellus and flagellum mediad triangular tooth at base of distal club-shaped lightbrown. part, distal part in inner-lateral view club-shaped with Thorax. Pronotum (Fig. 4) with central part of anterior a short dorsal tubercle at half of the length; inferiors and posterior lobe, median and lateral lobes (except in ventral view with triangular basal part one-third ventralmost part) brownish black, rest of anterior and of length; caudal part with basal half straight and posterior lobes castaneous; posterior lobe with caudal cylindrical, top constricted, bent in semicircle 90˚ processes rather wide apart, base solid, tapering, the dorso-axiad.

142 Chapter 5 Platystictidae of the Philippines

Measurements. Abdomen including appendages 29 1600-1800 m, 12-25.iii.1994 (Alex Buenafe) 1 female; North (29-30) mm, hind wing 19 (19-20) mm. Cotabato, Mt. Apo, Marbel river, 700-900 m, 12-25.ix.1994 Female. – As the male, but posterior margin of (Alex Buenafe) 1 male; Davao del Sur, Mt Talomo, Malagos, Baguio, Eagle Camp, Kal-lay Creek, 700-1000 m, x.1994 posterior lobe of pronotum without projections. (Alex Buenafe) 1 female; North Cotabato. Mt. Apo. Lake Measurements. Abdomen 27-28 mm, hind wing 20 Agko. 1200-1300 m, 29.iii. -2.iv.1995 (R. A. Müller) 5 males mm. 4 females; North Cotabato, Mt. Apo, Ilomavis, Lake Agko, Etymology. – Acutus (Latin), sharp; for the acute tooth Dum Creek. 1100-1200 m, ix.1995 (Alex Buenafe) 7 males on superior appendage. An adjective. 1 female; North Cotabato, Mt. Apo, Ilomavis, Kal-ay Creek, Sitio Sayaban. 1200-1300 m, ix.1995 (A. Buenafe) 3 males; Distribution (Fig. 99). – Philippine Islands: Luzon North Cotabato, Mt. Apo, Ilomavis, Lake Agko, Babang (Camarines Sur). Creek. 1600-1700 m, ix.1995 (A. Buenafe) 1 female.

Diagnosis. – Dark and medium-sized Drepanosticta; Drepanosticta aries Needham & Gyger differs from other species in the aries group by the (Figs 7-9, 99) robust superior appendages, i.e. approximately two Drepanosticta aries Needham & Gyger, 1941: 144-145, figs times as long as wide in dorsal view (appr. three times 3, 4, 8, 9 (original description, type locality Mindanao, in D. krios sp. n.), and the complete dark segment 8 Mnt Apo, Galog river, in MCZC) [type MCZN 23831, not examined]. – Hämäläinen & Müller 1997: 257, 276 of the abdomen. Superior appendages robust in lateral (distribution Mindanao). view, but those of D. acuta sp. n. (from Luzon) are significantly wider. Confined to Mt. Apo and Lake Material examined (all specimens Philippines, in RMNH). Sebu on Mindanao. – Mindanao. Tra’ankina. Lake Sebu. 21-23.xi.1991 (J. de los Reyes) 1 male; Tra’ankina. Lake Sebu. 3-5.i.1992 (J. de los Reyes) 1 female; Lake Sebu, Lamlahak, Talubek. 26.i. Male. – Head. Labrum, mandibles, anteclypeus dirty - 1.ii.1994 (L. Vinciguerra & E. Horn) 3 females; North white, narrow brownish black line along anterior Cotabato, Mt. Apo, Philipp. National Oil Comp. forest area, border of labrum and mandibles; rest of head bronze-

7

9 8

Figures 7-9. Drepanosticta aries Needham & Gyger, male [JvT 20089, Mindanao, North Cotabato, Mt Apo, Ilomavis, Lake Agko, 1100-1200 m, ix.1995]. – 7, pronotum, oblique view. – 8, anal appendages, dorsal view. – 9, idem, left lateral view. Scale bar 1 mm.

143 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

black; antenna with scapus and pedicellus yellowish on same sites) with lateral extremities of postorbital white, tip of pedicellus and flagellum brown; transverse carina acute rather than apiculate; pronotum with occipital carina well developed with lateral extremities posterior lobe indistinct, the erect lateral processes apiculate. absent, only with short erect collar. Thorax. Pronotum (Fig. 7) medium brown, but middle Measurements. Hind wing 22-24 mm, abdomen 31-32 of posterior lobe, and lateral lobe dark brown; anterior mm. lobe simple, but anterior ridge distinct; median and Status. – The name Drepanosticta aries Needham & lateral lobes without distinct features; posterior lobe Gyger is used here exclusively for specimens from with both sides approximately halfway a caudally Mount Apo, the type locality, and Lake Sebu, which is directed, flat and not erect process, approximately as situated c. 120 km south of Mt. Apo. Other specimens long as median length of posterior lobe of pronotum, with similar pronotum and appendages are known process ending in a 90˚ outward directed tip, somewhat from other places in Mindanao, from Eastern Samar, more robust than base, ca. one-fourth the length of Tawi Tawi and Sanga Sanga. They are assigned to base. Synthorax dark brown, except for a paler spot Drepanosticta krios sp. n. The other two species of this around spiracle. Legs pale, except for darker rings group occur in Luzon. on femora near tibiae. Wings hyaline, venation dark Distribution (Fig. 99). – Philippine Islands: Mindanao brown; Px 16-17 in fore wing, 15-16 in hind wing; (Mt. Apo, Lake Sebu) origin of R4+5 at subnodus, IR3 just distal to that level; Arculus arising just distal to Ax2, quadrangle in Drepanosticta krios van Tol, sp. n. forewing widening distally, anal veins shortly stalked; (Figs 10-12, 99) pterostigma castaneous, subquadrangular and oblique, proximal corner acute, distal side distinctly convex, Drepanosticta sp. n. 3. – Hämäläinen & Müller 1997: 257, cells between Costa and R1 distal to pterostigma rarely 276. divided. Type material. – Holotype male [JvT 18867] in RMNH: Abdomen. Dark brown, but paler basal annulae on ‘Philippines, Tawi Tawi / Languyan, 80-100 m / June 8, 1990 segments 3-7. Appendages (Figs 8-9) middle brown, / Roland A. Müller legit’. – Paratypes (all Philippine Islands, in RMNH, 112 males 22 females): Mindanao. Lanao del superiors base stout, in dorsal view tapering towards Norte, Kapatagan, Mt. Puting, Bato Sapad, 250 ft., x.1988 the tip, length ca. 2.5 times largest width, superiors (W. Catal) 10 males 1 female; Lanao del Norte, Sitio Tinago, actually fully flattened, thus hollow underside at base, 10-16.viii.1997 (A. Buenafe) 37 males 11 females; Lanao dorsally at three-fifths from base a very short dorsal del Norte, Iligan. Mimbalot Falls. 8˚10’50”N 124˚10’10”E. triangular tooth; inferiors in ventral view somewhat 29 m asl. 25.iii.2004 (V. Kalkman & J. van Tol) 14 males; Zamboanga del Norte Prov., 20 km S of Manucan, Labauan shorter than superiors, somewhat diverging, but tips Mts, 680 m, 15.x.1959 (Quate) 1 male; Zamboanga del directed inwards at c. 70˚. Norte Prov., Tampilisan, Gampoy river, 5-9.i.1991 (Th. Measurements. Hind wing 24-25 mm, abdomen Borromeo Jr) 3 males 1 female; Idem, Zamboanga del Sur, including appendages 35-36 mm, one specimen very Tigbao, Mt Timolan, Tigbao Creek, 12-21.viii.1994 (Th. small with hind wing 20 mm, abdomen including Borromeo) 28 males 4 females. – Sulu archipelago, Tawi appendages 30 mm. Tawi Id. Languyan, 80-100 m. 6-9.vi.1990 (R. A. Mueller) 15 males 3 females; Batu Batu, 14-16.iii.1991 (Th. Borromeo) 2 males; Sanga Sanga Id. 2.vi.1992 (Th. Borromeo) 1 male; Female. – Very similar to the male, but posterior lobe Tawi Tawi Id., Tarawakan. 100 m. 22-26.vi.1992 (Treadaway) of pronotum with lateral processes distinctly shorter, 1 male 2 females. without terminal knob, approximately the length of Excluded from type series. – Samar. Eastern Samar, Oras, 10- hind lobe and semi-erect; wings with anal veins sessile, 22.viii.1994 (Th. Borromeo leg) 1 male (locality doubtful). 17 Px in fore wing and hind wing. Some females Diagnosis. – A rather small Drepanosticta, similar in presumably conspecific with males of D. aries (collected structural details to D. aries Needham & Gyger, but

144 Chapter 5 Platystictidae of the Philippines

12 10

11

Figures 10-12. Drepanosticta krios sp. n., male [JvT 22092, Tawi Tawi, Languyan, 80-100 m, 9.vi.1990]. – 10, pronotum, oblique view. – 11, anal appendages, dorsal view. – 12, idem, left lateral view. Scale bar 1 mm. of smaller size, with flatter processes of the posterior pronotum, both ending in smoothly recurved, tapering lobe of the prothorax, the superior appendages much tip. Synthorax vivid brown, except for a paler spot more slender (especially visible in dorsal view), a lower around spiracle and a narrow, oblong spot along lower number of postnodal crossveins in both fore and margin of metepimeron. Legs pale, except for darker hind wings, and a conspicuous oblong bluish spot on rings on femora near tibiae. Wings hyaline, venation abdominal segment 8; differs from both D. acuta sp. n. brown; Px 14-15 in fore wing, 13-14 in hind wing; and D. rhamphis sp. n. in the smaller projections of the origin of R4+5 at subnodus, IR3 halfway first cell distal posterior lobe of the pronotum, and from the former to that level; Arculus arising at Ax2, quadrangle in in the straight rather than curved and tapered inferior forewing hardly widening posteriorly, anal veins shortly appendages, and from the latter in the dorsal tooth of stalked; pterostigma castaneous, width 1.6 times the the superior appendage. height, oblique, proximal corner acute, distal side distinctly convex, cells between Costa and R1 distal to Male (JvT 18867, holotype). – Head. Labrum, pterostigma rarely divided. mandibles, anteclypeus bluish white with a brownish Abdomen brown, but paler basal annulae on segments black line along anterior border of labrum and 3-7, segment 8 anteriorly with an oblong, bluish white mandibles, line on labrum in middle approximately spot, approximately two-fifths the length of segment, one-fifth the height of labrum, and tapering towards and posteriorly rounded. Appendages (Figs 11-12) the corners; rest of head bronze-black, coriaceous, with middle brown, superiors base stout, in dorsal view light-brown oval spots somewhat smaller than ocelli, tapering towards the tip, length ca. 3.5 times largest anterior to anterior ocellus and lateral to lateral ocelli; width, superiors flattened and curved, thus hollow antenna with scapus and pedicellus yellowish white, tip underside at base, dorsally at five-sevenths from base a of pedicellus and flagellum brown; transverse occipital very short dorsal triangular tooth; inferiors in ventral carina well developed with lateral extremities apiculate. view somewhat longer than superiors, parallel-sided, Thorax. Pronotum (Fig. 10) pale brown or dirty yellow, but tips directed inwards at c. 70˚. but middle of posterior lobe, and lateral lobe brown; Measurements. Hind wing 18.5 mm (18-19 mm); anterior lobe simple, with anterior ridge distinct; abdomen, including appendages 29 mm (29-30 mm). median and lateral lobe without distinct features; posterior lobe with both sides approximately halfway a Female. – Similar in coloration as male; structure of caudally directed, flat and not erect process, somewhat hind margin of posterior lobe more simple, straight, shorter than median length of posterior lobe of lateral parts ending sharp, below that level a broadly

145 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

rounded lobe; Px fore wing 16, hind wing 15; anal female of D. acuta sp. n. is distinctly different from the appendages brown, valves pointed and surpassing tip male; for differences with D. aries Needham & Gyger of cercus. and D. krios sp. n., see under these species. Measurements. Hind wing 19-20 mm; abdomen 28-29 Male [JvT 18991, holotype]. – Head. Labium brown; mm. labrum bluish white, anteriorly with narrow brown Note. – Dr. M. Hämäläinen confirms that the line, fading towards pale coloured part of labrum; specimen from Samar may be a case of mislabelling. mandibles bluish white with narrow brown anterior Most specimens collected by Mr Borromeo are line; anteclypeus bluish white, rest of head coriaceous, reliably labelled, but there are some other clear cases of black with metallic shine; transverse occipital carina mislabelling in the collection. Usually, mislabelling is well-developed, the lateral extremities acutely angulate. much higher in collections of commercial collectors. Thorax. Pronotum (Fig. 13) multi-coloured, with This is an opportunity to emphasize the professionalism middle part of anterior lobe, central parts of both of the collectors working for Roland A. Müller. halfs of median lobe, posterior part of lateral lobe, Etymology. – Krios (Greek), ram; for its resemblance to and central part of posterior lobe brownish black, D. aries Needham & Gyger. A noun in apposition. rest much paler; structure of anterior, median and Distribution (Fig. 99). – Philippine Islands: Mindanao, lateral lobes simple, posterior lobe relatively long, Tawi Tawi, Sanga Sanga. The locality of the specimen with a paired posterior process, long, ending abruptly from Samar is considered doubtful, and not on the in an antero-laterally directed part, turned 110-120˚ map. backward, the tip with long and stout setae. Synthorax completely brownish black, legs fully yellowish white. Wings hyaline, venation brown, lighter in basal half; Drepanosticta rhamphis van Tol, sp. n. Px 16 in fore wing, Px 16 in hind wing; R4+5 at or (Figs 13-15, 99) even just proximal to Ax2; IR3 arising halfway first Drepanosticta sp. n. 11. – Hämäläinen 1997: 257, 277 cell distal to that level; Arculus just distal to Ax2; (distribution Catantuanes I.). quadrangle somewhat widening distally in fore wing; anal veins asymmetrical, sessile or very shortly stalked; Type material. – Holotype male [JvT 18991] in RMNH: pterostigma oblique, width c. 1.7 times the height, ‘Philippines, Catanduannes [sic!] Id / Gigmoto, San Pedro / proximal side acutely angulate, distal side convex; cells Egwang Tapayas Creeks / June 20.-30. 1996, 300-500 m / beyond pterostigma between Costa and R1 undivided. Alex Buenafe legit / Coll. Roland A. Müller’. – Paratypes (all Abdomen. Middle brown with pale, bluish white, Philippine Islands, in RMNH): Catanduanes Id, Gigmoto, San Pedro, Simohe Creeks, 300-500 m, 20-30.vi.1996 (A. anterior markings on each segment as follows: segment Buenafe) 4 males; same, Tongao Creeks, 400-500 m, 20-30. 2 narrow, covering anterior 2/5th of segment, segments vi.1996 (A. Buenafe) 1 female. 3-6 with dorso-anterior spot of 1/10th of segment Non-type material. – Luzon Id, Camarines Norte, Sn Lorenzo length, segment 7 pale in anterior 1/6th of segment, Ruiz, San Isidro, Sitio Bakalid, Patag river, 300-500 m, 26-27. rest of anterior half of this segment lightbrown, ix.1997 (C.M. Nazareno) 1 female. segment 9 with small bluish anterior spot, segment 10 pale bluish (?). Appendages (Figs 14-15) creamish; Diagnosis. – A medium-sized species with dark superiors in dorsal view with stout base, approximately synthorax and paler brown abdomen with distinct 2/3 of segment length ending in small dorsal tooth, small bluish markings anteriorly on abdominal distal part scoop-like and vertically oriented; inferiors segments 3-6; hind margin of hind lobe of pronotum rather slender, distal half bent outwards, the tip with a pair of hammer-like projections. The female strongly tapered and first bent axiad, then dorsad. of D. rhamphis sp. n. has similar projections on the Measurements. Hind wing 19 mm, abdomen including posterior lobe of the pronotum as the male, while the appendages 28.5 mm.

146 Chapter 5 Platystictidae of the Philippines

14

13

15

Figures 13-15. Drepanosticta rhamphis sp. n., male [JvT 18858, Catanduannes, Gigmoto, San Pedro, Simohe Creeks, 20-30. vi.1996]. – 13, pronotum, oblique view. – 14, anal appendages, dorsal view. – 15, idem, left lateral view. Scale bar 1 mm.

Female. – As the male, but posterior processes of species; inferiors in ventral view more or less clasper- posterior lobe of pronotum somewhat shorter, of like, the tip curved medio-dorsad. similar shape as in the male. Presumably the sister-group of the D. lymetta + D. Measurements. Hind wing 19 mm, abdomen 28 mm. megametta group. Included species: D. belyshevi Hämäläinen, D. Etymology. – Rhamphis (Greek): hook; after the flavomaculata sp. n. , D. trachelocele sp. n. structure of the processes at hind margin of posterior Distribution: Bohol, Leyte, Panaon, Samar, Mindanao, lobe of pronotum. A noun in apposition. Camiguin Id. Distribution (Fig. 99). – Philippine Islands (Luzon region): Catanduanes, Luzon. Drepanosticta belyshevi Hämäläinen (Figs 16-18, 100) Drepanosticta belyshevi group Drepanosticta belyshevi Hämäläinen, 1991: 65-68, figs 1-4 Transverse occipital carina distinct, with lateral (original description, holotype male Philippines, Panaon extremities sharp, but not apiculate; lateral corners Id., San Francisco, Anislagon Gamay (200 ft), Aug 1988, W. Catal leg. [in Müller collection, now in RMNH] of anterior lobe or pronotum distinctly widened or [examined], distribution Panaon Id). – Hämäläinen even with a long process; posterior lobe with a pair of & Müller 1997: 257, 276 (distribution East Visayan rather short, flat, curved or folded processes; synthorax subregion: Leyte, Panaon, Bohol). with characteristic pale markings in posterior corner Material examined (all Philippine Islands, in RMNH). – of metepisternum and metepimeron; pterostigma Bohol, Sierra Bullones, Pilar, 10-11.iv.1989 (W. Catal) 4 somewhat wider than high; superior appendages with males. – Samar, Samar Prov., Hinubangan, San Isidro, San ventral side of base (lateral view) curved sharp dorsad, a Isidro river, 90-200 m, 31.iii. - 5iv.1997 (R.A. Müller) 1 sharp ventral tooth near base, but size variable between female. – Leyte, S. Leyte, Mahaplag, Hilusag, Mt Balocaue,

147 18 16 17

Figures 16-18. Drepanosticta belyshevi Hämäläinen, male [JvT 22216, Bohol, Sierra Bullones, Pilar, 10-11.iv.1989]. – 16, pronotum, oblique view. – 17, anal appendages, dorsal view. – 18, idem, left lateral view. Scale bar 1 mm.

700 m, 30.xi. - 2.xii.1989 (Th. Borromeo jr) 2 males 1 female; Palawan has an extensive pale stripe over posterior idem, Magsuganao river, 18-30.ix.1990 (Th. Borromeo jr) three-quarters of metepisternum and very distinct 2 males. – Panaon, Southern Leyte prov., San Francisco. superior appendages; Protosticta lepteca sp. n. can be Mt Kaneo, 400 ft, viii.1988 (W. Catal) 2 males 1 female; distinguished by the absence of the Ab vein, and the idem, San Francisco, Anislagon Gamay, 200 ft, viii.1988 (W. Catal) 1 male (holotype) 1 female; idem, San Francisco, structure of dorsal appendages, which are not clasper- Panan-awan Creek, viii.1988 (W. Catal) 1 male; idem, San like. D. flavomaculata sp. n. is considered to be closely Francisco, Batong Lapad, viii.1988 (W. Catal) 1 male; idem, allied, but lacks the projections of the anterior lobe of San Francisco, Gabing Gamay, big river, x.1988 (W. Catal) the pronotum. 1 male; idem, San Francisco, Tabon river Lilo-an, x.1988 (W. Catal) 1 male; idem, San Francisco, Mt Anislagon, 350 ft, viii.1988 (W. Catal) 2 male; idem, Anislagon river, 10- Male. – Head. Labium brown, labrum and anteclypeus 12.x.1990 (Th. Borromeo jr) 3 males 3 females. bluish white, labrum with apical third brown, mandibles pale, bluish, but somewhat darker than Diagnosis. – A relatively slender and pale brown labrum; rest of head brownish black, frons and vertex coloured Drepanosticta, with conspicuous processes coriaceous; transverse occipital carina distinct, lateral on the sides of both the anterior and the posterior extremities sharp, but not apiculate. Antenna with lobe of the pronotum, and posterior one-third of scapus and pedicellus yellowish white, flagellum brown. metasternum and metepimeron bluish or yellowish Thorax. Pronotum (Fig. 16) pale brown, the median white. Shares remarkable structure of anterior lobe of lobe pale yellowish, anterior lobe laterally with a pair pronotum with D. trachelocele sp. n., D. moorei Van of dorsally erect cylindrical processes, the tip directed Tol & Müller, D. quadricornu sp. n. and Protosticta outward, length in lateral view nearly as high as lateral lepteca sp. n. The projected corners of the anterior lobe; median and lateral lobes simple; hind margin of lobe of the pronotum, although only known for posterior lobe medially acute, laterally with a paired species in the Philippines, is considered a parallel flat process, in lateral view approximately as long as evolution, and not a synapomorphy. Its closest median lobe, directed posteriad, curved outward, relative is Drepanosticta trachelocele sp. n. from Samar with a short triangular spine as base. Synthorax (for differences see under that species); D. moorei medium brown, but the following parts yellowish: can immediately be distinguished on the presence of small marking of mesepimeron, approximately one- a broad pale band over metepisternum and dorso- third of metepisternum posterior to metastigma, and posterior third of metepimeron; D. quadricornu from approximately one-third of metepimeron. Wings

148 hyaline, venation brown, Px 15-16 in fore wing, Px 14- Occurrence. – Apparently occurring only in low 15 in hind wing; R4+5 arises well distal to subnodus, densities on most sites, since only short series are IR3 approximately halfway first cell distal to that level; available. Rather widespread, but specimens from Arculus arises distal to Ax2; quadrangle somewhat Samar are here considered specifically distinct, widening in fore wing, scarcely so in hind wing; anal Drepanosticta trachelocele sp. n. veins asymmetrical and shortly stalked, pterostigma Distribution (Fig. 100). – Philippine Islands (East width c. 1.9 times the height, brown with narrow Visayan subregion): Leyte, Panaon and Bohol. pale line against veins; some cells distal to pterostigma between Costa and R1 divided. Drepanosticta flavomaculata van Tol, sp. n. Abdomen. First segments medium brown, more caudal (Figs 19-21, 100) segments darker, especially segment 3-6 with yellow Drepanosticta sp. n. 18. – Hämäläinen & Müller 1997: 258, basal annulae. Appendages (Figs 17-18) with superiors 277 [Mindanao, Camiguin]. in dorsal view basal half very stout, the caudal half Type material. – Holotype male [JvT 20359] in RMNH: slender and dorso-ventrally flattened, bent as a forceps, ‘Philippines. Mindanao Id / Bukidnon, Kalatungan Mts / the tip squarish; inferiors in ventral view somewhat Pangantocan, Brgy Mendes / Mikaramagan Creek, Mandom longer than superiors, nearly straight, the tip bent / June 1995. 900-1200 m / Alex Buenafe legit’ . – Paratypes 90˚dorsad. (all Philippine Islands, in RMNH, total 160 specimens) (by Measurements. Hind wing 18-20 mm, abdomen island in chronological order): Camiguin Id. Mambajao. including appendages 27-32 mm. Brgy Pandan, Mt Timpo-ong, Katibawasan Falls, 500-700 m, 22.v.1995 (A. Buenafe) 6 males 1 female; Catarman, Tuasan Falls, 29.x.2003 (R.J. Villanueva) 2 males. – Mindanao. Female. – As the male, but abdomen distinctly stouter; Zambo[a]nga d. Norte. Manucan, 20 km So. Labauan Mts, structure of prothorax as male, but processes of anterior 680 m, primary forest, 15.x.[19]59 (Quate) 1 male; South and posterior lobe of prothorax shorter and more Cotabato, Salacafe, El Milil, 1250 m, 7.iv.1985 (R.A. Müller) slender; last abdominal segments stout, brown, the 2 males; [South Cotabato, Koronadal], Barrio 8, 13.iv.1985 (R.A. Müller) 2 males; South Cotabato, Koronadal, Barrio valve relatively slender, the stylus not surpassing the 8, 100-200 m, 12-14.vii.1986 (R.A. Müller) 11 males 5 cercus. females; South Cotabato, Koronadal, Barrio 8, 19.vii.1987 (J. Mesurements. – Hind wing 20-22 mm, abdomen 28- de los Reyes) 1 male; South Cotabato, Koronadal, Barrio 8, 32 mm. 30.vi.1991 (J. de los Reyes) 1 male 1 female; North Cotabato,

19 21 20

Figures 19-21. Drepanosticta flavomaculata sp. n., male [JvT 20225, Mindanao, Koronadal, Barrio 8, 4.ii.1994]. – 19, pronotum, oblique view. – 20, anal appendages, dorsal view. – 21, idem, left lateral view. Scale bar 1 mm.

149 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

Alamada, Mt Makatoring, 700-900 m, 1000 m, 1.ii.1992 Male [JvT 20359, holotype]. – Head. Labrum, (Th. Borromeo) 10 males; South Cotabato, Koronadal, Barrio mandibles, and anteclypeus ivory white, anterior one- 8, 29.viii.1993 (J. de los Reyes) 5 males; South Cotabato, fifth of labrum and mandibles brownish black; rest Mt Matutum, 500-700 m, 16-19.ix.1993 (Th. Borromeo) 2 males; South Cotabato, Koronadal, Barrio 8, 4.ii.1994 (L. of head bronze-black, frons and vertex coriaceous; Vinciguerra & E. Horn) 2 males; Davao del Sur, Malagos, transverse occipital carina distinct, lateral extremities Baguio, Eagle Camp, 500-600 m, 28.iii.1995 (R.A. Müller) 1 sharp, not apiculate. Antenna with scapus and male; Davao Oriental, Baganga, Brgy Upper Mikit, Mangoy pedicellus dirty yellow, flagellum brown, somewhat Falls, 300-350 m, 6.iv.1995 (A. Buenafe & A. Gorostiza) paler at base. 1 female; Davao Oriental, Cateel, Aliwagwag Falls, 200- Thorax. Pronotum (Fig. 19) with anterior lobe 300 m, 10.iv.1995 (A. Buenafe & A. Gorostiza) 1 male 1 female; Data as holotype, 5 males 1 female; Bukidnon, Mt castaneous, dull, median and lateral lobes dirty yellow, Kalatungan, Talakag, Brgy Mebadiang, Magamanason Creek, only dorsal parts somewhat obscured; posterior lobe Sitio Olayan, 1000-1100 m, 5-18.viii.1995 (A. Buenafe) 4 brown; anterior lobe erect, lateral sides broad and males 6 females; Bukidnon, Mt Kalatungan, Pangantocan, flat, median and lateral lobes simple; posterior lobe Sitio Mandon, Mekaramagan Creek, 1800 m, 12-15.viii.1995 with lateral portion on innerside with flat, short, (A. Buenafe) 4 males; Bukidnon, Mt Katanglad, Impasugong, Brgy Impulatao, Gantongan Creek, 800-900 m, 19-29. recurved process, just somewhat longer than length viii.1995 (A. Buenafe) 9 males; Bukidnon, Mebadiang, of median portion of posterior lobe. Synthorax Dumatap, Mt Kalatungan, Mansabilan Creek, 1000-1300 castaneous, coriaceous except for metepimeron and m, 24.xi.1995 (A. Buenafe) 3 males 1 female; Bukidnon, metepisternum, with a squarish pale marking just Mebadiang, Dumatap, Mt Kalatungan, Muntian Creek, before posterior margin of metepisternum, and a 1200-1400 m, 24-25.xi.1995 (A. Buenafe) 6 males 2 females; similar marking in posterior corner of metepimeron. Surigao del Sur, Tabon, Tabon Falls, 100-200 m, 26.v. -8.vi.1996 (A. Gorostiza & A. Buenafe) 9 males 3 females; Wings hyaline, venation brown; Px 16 in fore wing, Davao Oriental, Boston, Mt Agtuuganon, Camp 55, 1020 PX 15 in hind wing; R4+5 arising at or just distal to m, 29.v. -7.vi.1996 (R.A. Müller et al.) 4 males 4 females; subnodus; IR3 circa halfway first cell distal to that Davao Oriental, Sigaboy, Tandang Sora, Yakal, Tabamban level; Arculus arising distinctly distal to Ax2 in fore River, Lakahan Creek, 400-800 m, 5-14.x.1996 (A. Buenafe) wing, c. distance between R+M and CuP; quadrangle 7 males 3 females; Davao Oriental, Sigaboy, Tandang Sora, Yakal, Tabamban River, Buyo Creek / Tubonol Creek / widening in fore wing; anal vein long and symmetrical; Langanisan Creek, 400-800 m, 5-14.x.1996 (A. Buenafe) 3 pterostigma width c. 1.5 times the height, a narrow males; Lanao del Norte, Linamon, Sitio Tinago, 200-400 m, pale border against veins; many cells between Costa 10-16.viii.1997 (A. Buenafe) 15 males 6 females; Bukidnon, and R1 distal to pterostigma divided. Katanglad Mts, Impasugong, Impalutao, Gantongan Creek, Abdomen. Segment 1 brown, each more caudal 1200-1400 m, 17-28.viii.1997 (A. Buenafe) 4 males; Davao segment darker, except for segment 10, which is City, Datu Salumay, 25 Feb 2004 (R.J. Villanueva) 1 males; Iligan, Tinago Falls, 8˚09’33”N 124˚11’11”E, 160 m, completely blue dorsally; segments 3-7 with basal one- 25.iii.2004 (V. Kalkman & J. van Tol) 4 males. sixth to one-eighth with yellow marking. Appendages (Figs 20-21) dirty yellow, superiors with stout base, Diagnosis. – Based on the structure of the anal distal three-fifths in dorsal view clasper-like, tip appendages, and the coloration of the synthorax, in one squarish, in lateral view distal half distinctly flat, a group with D. belyshevi Hämäläinen. Distinguishable sharp, but rather inconspicuous, ventro-mediad tooth from other species in this group by the absence of a near base; inferiors approximately as long as superiors, conspicuous, paired process on the anterior lobe of stout at base, distal three-fifths approximately the pronotum, the curved and flat processes on the cylindrical, bent 80º inward, innerside emarginate, tip posterior margin of the posterior lobe of the pronotum, directed dorsad (caudal view). and a poorly developed ventro-medial tooth on the Measurements. Hind wing 24 (20-24) mm, abdomen superior appendage of the male. (including appendages) 34 (30-34) mm.

150 Chapter 5 Platystictidae of the Philippines

24 23 22

Figures 22-24. Drepanosticta trachelocele sp. n., male [JvT 20331, Samar, Hinabangan, San Isidro, 100-200 m, 29.viii-20. ix.1996]. – 22, pronotum, oblique view. – 23, anal appendages, dorsal view. – 24, idem, left lateral view. Scale bar 1 mm.

Female. – Coloration as the male, but structure of Western Samar province, San Jose de Buan, Mt Hurao, San pronotum distinctly different, with processes on Andres. 14-20.v.1997 (A. Buenafe) 2 males. posterior lobe short and erect, flat, although with much variation in length and shape between specimens. Diagnosis. – One of the very few species of Innerside of these processes rounded, outerside sharply Drepanosticta with paired processes on hind margin projected in some specimens, broadly rounded in of both the anterior and the posterior lobe of the others, but process may even be shaped as a small pronotum; most closely related to D. belyshevi triangular, erect projection. Hämäläinen, from which it differs particularly by the Measurements. Hind wing 22-23 mm, abdomen 30-32 structure of the processes of the posterior lobe of the mm. pronotum: the processes in D. belyshevi are placed close to the median line and curved over the abdomen, Etymology. – Flavomaculatus (Latin), with yellow whereas the processes in D. trachelocele are placed near markings; for the coloration of the synthorax. An the lateral corners, with a short erect base and the rest adjective. recurved ventrad. For differences with other Philippine Distribution (Fig. 100). – Philippine Islands Drepanosticta species with projections on anterior lobe (Mindanao subregion): Camiguin Island, Mindanao. of pronotum, see D. belyshevi Hämäläinen. Rather widespread in Eastern Mindanao region. Male [JvT 20323, holotype]. – Head. Labium, mandibles and anteclypeus bluish white, a brownish Drepanostocta trachelocele van Tol, sp. n. black margin on anterior one-fifth of labium and one- (Figs 22-24, 100) eighth of mandibles; rest of head brownish black with Type material. – Holotype male [JvT 20323] in RMNH: bronze metallic shine, frons and vertex coriaceous; ‘Philippines, Samar Id / Samar Prov., Hinubangan / Arizona, transverse occipital carina well developed, with acute 100-280 m / March 29.-April 6. 1997 / R.A. Müller leg. et coll.’, ‘Drepanostocta belyshevi Hämäl. (ssp. ?) M. Hämäläinen lateral extremities, more angulate that in D. belyshevi; det.’. – Paratypes (all Philippine Islands, in RMNH, 18 antenna with scapus and pedicellus dirty yellow; specimens): Samar Id, Hinabangan, Bagakay, Arisona, 100- flagellum castaneous. 200 m, 29.viii-20.ix.1996 (A. Buenafe) 4 males 1 female; Thorax. Pronotum (Fig. 22) with anterior lobe same site, but San Isidro, 100-200 m, 100-200 m, 29.viii-20. castaneous, somewhat darker at hind margin, middle ix.1996 (A. Buenafe) 1 male 3 females; same site as holotype, 3 males 1 female; same site, but San Isidro, San Isidro river, and lateral lobes dirty yellow, posterior lobe pale brown 90-200 m, 31.iii.-5.iv.1997 (R.A. Müller) 1 male 2 females; with a rounded or sub-triangularly shaped marking in

151 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

middle, hind margin of posterior lobe also somewhat this species is considered the sister-species of D. darker; anterior lobe with a paired process, laterally on belyshevi Hämäläinen. lobe, rounded and directed anterio-dorsad, total length Etymology. – Trachelocele: a genus of antelopes; for the surpassing median length of pronotum; hind margin of shape of the processes on the pronotum. A noun in posterior lobe with a paired process, distance between apposition. bases of processes more than five times the width of Distribution (Fig. 100). – Philippine Islands: Samar. process at base, process halfway curved backward (ventrad) and inconspicuous in many specimens (much D. moorei group variation), but not a long, flat and curved process as in D. belyshevi. Synthorax castaneous, but meso- and Transverse occipital carina poorly developed, lateral metakatepisternum somewhat paler, a subsquarish extremities absent or just discernable; pronotum with bluish white spot posteriorly on metepisternum, and processes on anterior and posterior lobe distinctly a similar spot on metepimeron, the latter one with a different between species; synthorax with long and ventro-lateral projection. Legs dirty yellow, obscure distinct stripes over mesepisternum and mesepimeron; brown rings on two-thirds the length of femora. Wings pterostigma slightly wider than high; superiors strongly hyaline, venation brown; Px 16 in fore wing 16, Px 15 flattened dorso-ventrally, and broadly triangular in hind wing; R4+5 distal to subnodus, IR3 halfway in lateral view; base of inferiors in ventral view first cell distal to subnodus; Arculus distal to Ax2; subparallel, the top as a clasper, with a more or less Ab vein joining Ac just before wing margin (Y-veins distinct ridge on inner margin; tip sharp. subsessile); pterostigma castaneous, oblong, c. two Based on structure of male anal appendages and the times wider than high; veins distal to pterostigma coloration of the synthorax, the species of this group numerous, but undivided. are not members of the group of ‘typical Philippine Abdomen castaneous, with creamish or bluish white Drepanosticta’ species. Their affinities are presently markings as follows: segments 3-6 with a narrow ring unsettled. against anterior border of segment, the white marking Included species: D. luzonica sp. n., D. moorei Van Tol projecting posteriorly on both sides of median line & Müller. of tergites against borders; segment 8 with yellowish Distribution: Luzon. white anterior ring, c. one-fourth of length of segment. Appendages (Figs 23-24) very pale brown, superiors dorsally stoutly built, distal half as a clasper in dorsal Drepanosticta luzonica van Tol, sp. n. view, and club-shaped on innerside; inferiors in ventral (Figs 25-27, 101) view very slender, straight, the top bent medio-dorsad. Drepanosticta sp. n. 9. – Hämäläinen & Müller 1997: 257, Measurements. Hind wing 19 (18-19) mm; abdomen 277. including appendages 30 (28-32) mm. Type material. – Holotype male [JvT 9451] in RMNH: ‘Philippines / [Luzon] Ifugao Province / Jacmal Bunhian / 24 Female. – Similar to male, including structural details km E Mayoyao 800- / 1000m 7-8.iv.1967’. of the processes of the anterior and posterior lobes of the pronotum, although both pairs of processes are Diagnosis. – A relatively large species, with somewhat smaller, especially the ones on the anterior conspicuous yellow markings on synthorax. Based lobe. on synthoracic pattern, superficially resembling Measurements. Hind wing 18-20 mm, abdomen 27-29 Drepanosticta lestoides Brauer, but superior anal mm. appendages distinctly different, lacking the superior tooth, and the tip squarish rather than smoothly Affinities. – Based on the structure of the pronotum, rounded. Differs from D. moorei Van Tol & Müller by

152 Chapter 5 Platystictidae of the Philippines

27 25 26

Figures 25-27. Drepanosticta luzonica sp. n., male [JvT 9451, holotype]. – 25, pronotum, oblique view. – 26, anal appendages, dorsal view. – 27, idem, left lateral view. Scale bar 1mm.

the absence of conspicuous processes on anterior and Wings hyaline, venation brown, basal half paler; Px posterior lobes of pronotum. 13 in fore wing, Px 12 in hind wing; Arculus arising at Ax2; R4+5 arising at subnodus; IR3 halfway cell Male [JvT 9451]. – Labrum, anteclypeus and mandibles distal to R4+5; quadrangle oblong and quadrangular, bluish white, partly yellowish white; anterior one-fourth not widening posteriorly; pterostigma rhomboid, ca of mandibles brownish black; rest of head bronze-black, 1.5 times as wide as high; cells between Costa and R1 frons and vertex coriaceous; postoccipital carina without beyond pterostigma undivided. extremities; antenna with scapus brownish black, Abdomen. Brown, lower parts of segments 1-2 pedicellus brown, flagellum broken in type. yellowish white, anterio-basal part of segments Thorax. Pronotum (Fig. 25) dark with distinct, pale 3-7 pale (partly discoloured in type), but dark line yellowish white markings; anterior and median lobe extending over full length; lower half of segments 8-9 yellowish white with higher parts of median lobe pale; appendages (Figs 26-27) brown, superior club- brown; posterior lobe with median portion brownish shaped distal part with tip squarish, the ventral margin black, lateral portions greyish brown, lateral lobes forming a broad triangular tooth; superiors without brownish black, but central parts with pale spots; dorsal tooth; inferiors in distal half very slender, curved anterior, median and lateral lobes simple; posterior lobe inward, the very tip dorsad, lacking an inward directed with lateral portions semi-erect, somewhat extended tooth. ventrad, broadly rounded caudally. Synthorax bronze- Measurements. Hind wing 23 mm, abdomen including black with pale yellow markings as follows: dorsal appendages 38 mm. carina, a small and narrow line dorso-posteriorly on mesepisternum; metepisternum nearly fully pale, Female. – Unknown. except for posteriormost part, anteriorly extending over lower part of mesepimeron and mesokatepisternum; Etymology. – Luzonicus (Latin), from Luzon, for the lower half of metepimeron, pale marking extending type locality. An adjective. posteriad and covering the full width in posterior quarter of metepimeron. Distribution (Fig. 101). – Philippine Islands: Luzon

153 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

28 30 29

Figures 28-30. Drepanosticta moorei Van Tol & Müller, male [JvT 18624, Luzon, Nueva Viscaya, Sta Fe, Atbo river, 550-800 m, 10.vi.1991]. – 28, pronotum, oblique view. – 29, anal appendages, dorsal view. – 30, idem, left lateral view. Scale bar 1 mm.

(central northern part). Apparently a local or rare and the conspicuous greenish yellow stripe over species; not represented in the Müller collection. synthorax. For a comparison of species with long projections of the anterior lobe, see D. belyshevi Hämäläinen; for a comparison with other species with Drepanosticta moorei Van Tol & Müller pale stripe over synthorax, see D. luzonica sp. n. (Figs 28-30, 101)

Male [holotype, JvT 22225]. – Head. Labrum Drepanosticta moorei Van Tol & Müller, 2003: 42 [original and anteclypeus very clear blue, anterior border of description, holotype male Luzon, Nueva Viscaya, Sta Fe, Atbo River, 550-800 m, 10 Jun 1991, R.A. Müller] labrum with narrow black line; mandibles brown, but [examined]. inner basal corner blue; rest of head bronze-black; Platysticta annulata Selys, 1886: 156 [partim]. antenna with scapus and pedicellus dirty yellow, the Drepanosticta sp. – Lieftinck 1961: 136 [specimen ‘588’ in tip of pedicellus somewhat darker, flagellum brown; Selys collection, IRSN Brussels, a paralectotype of D. transverse occipital carina indistinct and lacking annulata (Selys), not conspecific with lectotype]. conspicuous extremities. Drepanosticta sp. n. 7. – Hämäläinen & Müller 1997: 257, 276-277 [distribution Luzon]. Thorax. – Pronotum (Fig. 28) with anterior and median lobe yellowish, lateral lobe brown, posterior Material examined [all Philippine Islands, in RMNH]. – lobe dirty yellow; anterior lobe approximately halfway Luzon. Nueva Viscaya, Dalton Pass, 850-900 m, 8-9.vi.1991 each side with very long subcylindrical process, (R.A. Müller) 1 male; Nueva Viscaya, Sta Fe, Atbo river, touching median lobe, and reaching virtually hind 550-800 m, 10.vi.1991 (R.A. Müller) 12 males 10 females margin of central portion of posterior lobe; median and [including holotype]; Nueva Ecija, Caranglab, Batching river, 700-850 m, 11.vi.1991 (R.A. Müller) 2 males; Aurora lateral lobes simple; hind lobe laterally with enormous Prov., Dilalongan, Sitio Biyak, Dibaraybay, 300-500 m, collar, half-round, nearly twice as long as basal part of 3-15.vii.1996 (C.M. Nazareno) 1 female; Zambales Prov., posterior lobe; lateral margin with long and thin setae. Masinloc, Mt Coto, Tal-tal, 400-700 m, 1-8.iv.1997 (C.M. Synthorax bronze-black, but metepisternum nearly Nazareno) 1 female. fully greenish yellow, and dorso-posterior third of metepimeron greenish yellow. Legs dirty yellow. Wings Diagnosis. – Unmistakable species by the combination hyaline, venation brown; Px 13 in fore wing, Px 13 of long processes on the anterior lobe of the pronotum, in hind wing; origin of R4+5 just distal to subnodus,

154 Chapter 5 Platystictidae of the Philippines

IR3 arises six cells distal to nodus in fore wing, five D. lymetta group cells in hind wing; Arculus just distal to Ax2, distinctly stalked; Ac and Ab forming a Y, subsessile; number Transverse occipital carina distinct, with acutely of cells between distal end of quadrangle and place angulate lateral extremities; anterior lobe of pronotum where CuP meets hind margin in fore wing 5, in hind simple, without projections or processes; posterior wing 7; pterostigma brown, with a narrow line against lobe of pronotum with a pair of processes on hind veins; ca. 1.6 times longer than high, the proximal side margin, typically divided or hooked at top; synthorax oblique, the distal side convex; cells between Costa and castaneous; pterostigma distinctly wider than high; R1 posterior to pterostigma undivided. superior appendages with a blunt dorsal tooth and a Abdomen brown, but basal one-tenth of segment sharp medially directed one on inner surface; inferior 3, basal one-sixth of segments 4-6, basal one-third appendages slender and straight, the tip emarginate, of segment 7 dirty yellow; segments 8-10 tergites curved 90˚ medio-dorsad, undivided. Based on the brown. Appendages (Figs 29-30) greyish brown, top structure of the anal appendages, presumably the sister- of inferiors castaneous; superiors with basal one-fourth group of the D. megametta group. stout, rest narrow in dorsal view, actually strongly Included species: D. clados sp. n., D. furcata sp. n., D. flattened dorso-ventrally, and broadly triangular in hermes sp. n., D. lymetta Cowley, D. taurus Needham lateral view; inferiors in ventral view subparallel, the & Gyger. top bent outward, then inward in a semicircle, in Distribution: Mindanao and Siquijor. lateral view the top bends in 90˚. Measurements. Hind wing 22 (20-22) mm; abdomen Drepanosticta clados van Tol, sp. n. including appendages 37 (31-37) mm. (Figs 31-34, 102)

Female. – Very similar to male, including coloration Type material. – Holotype male [JvT 26817] in RMNH: and structure of pronotum; segments 8-10 brown, anal ‘[Philippine Islands, Mindanao] 002-XII-7-2003 / Spring appendage light brown, genital valves surpassing apex -Malabog, Pakibato Dist., Davao City / R.J. [Villanueva]’. – of appendages. Paratype: 1 male, same data [in RMNH]. Measurements. Hind wing 21-22 mm, abdomen 30-31 Diagnosis. – An inconspicuous brown species; mm. posterior processes of hind lobe of pronotum extremely long, c. 80% of median length of pronotum, the tip Affinities. – Van Tol & Müller (2003: 44) assumed bifid, both branches of equal length; also characterised a close relationship of this species with D. belyshevi in the lymetta-group by its long inferior appendages, Hämäläinen, particularly based on the structure of distinctly surpassing the hooked superiors in lateral the processes of the anterior lobe of the pronotum. A view (Fig. 33). more recent analysis revealed that the structure of the pronotum of both species is distinctly different, and the Male [holotype, JvT 26817]. – Head. Labrum, projections cannot be considered homologous. Based anteclypeus and mandibles ivory or bluish white, on the coloration of the synthorax and the structure anterior one-third of labrum and mandibles brown; of the anal appendages of the male, D. moorei must be rest of head bronze-black; transverse occipital carina considered the sister-species of D. luzonica sp. n. distinct, with distinctly angulate lateral extremities; Distribution (Fig. 101). – Philippine Islands: Central antenna with scapus brownish-black, pedicellus brown, Luzon (provinces Nueva Viscaya, Nueva Ecija, Aurora flagellum unknown (missing). and Zambales). Apparently occurring in low densities Thorax. Pronotum (Figs 31-32) rather dark, with over a relatively large area; in larger numbers on type anterior lobe wide, but relatively simple with only locality only. anterior margin distinctly erect, brown in median part,

155 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

32

33

31

34

Figures 31-34. Drepanosticta clados sp. n., male [JvT 26816, Mindanao, Davao City, Pakibato District, Malabog, 7.xii.2003]. – 31, pronotum, oblique view. – 32, pronotum, frontal view of projection of hind lobe. – 33, anal appendages, dorsal view. – 34, idem, left lateral view. Scale bar 1 mm. paler in lateral corners; median lobe with a paired, one-sixth of segment length, one-seventh on segments convex, brownish black protuberance, better defined 3-4, one-eighth on segments 5-6, and one-seventh than in D. lymetta; posterior lobe castaneous, medially on segment 7; segments 8-10 brown, but segment somewhat darker, posteriorly with a paired long 10 presumably paler in life. Appendages (Figs 33-34) process, c. 80% of length of pronotum itself, apically yellow; superiors in lateral view distinctly hooked, basal V-shaped, the branch of equal length and stoutly third directed dorsad, distal two-thirds slightly ventrad; built. Synthorax castaneous, but mesokatepisternum, base of terminal part with short, triangular tooth, and metakatepisternum, metepimeron and anterior part just distal to that a similar distal tooth; terminal two- of metepisternum darker, no pale markings. Legs thirds flattened, the tip squarish; inferiors in ventral with fore coxa and upper part of middle coxa dark view straight, the terminal one-sixth more slender, brown, rest of legs yellowish or reddish brown. Wings tapering and curved dorso-mediad. hyaline, veins brown; Px 15-16 in fore wing, Px 15 in Measurements. Hind wing 21.5 mm; abdomen hind wing; R4+5 arising from subnodus, IR3 arising including appendages 33 mm. half a cell distal to that level; Arculus just distal to Female. – Unknown. Ax2; Y-shaped anal veins virtually sessile, distinctly asymmetrical; pterostigma 1.8 times as wide as high, Etymology. – Clados: branch (Greek), for the structure innerside angulate; cells between Costa and R1 distal of the processes of the posterior lobe of the pronotum. to pterostigma undivided. A noun in apposition. Abdomen. Brown, with pale hastate markings dorsally Distribution (Fig. 102). – Philippine Islands: Eastern on segment bases as follows: segment 2 very small, Mindanao, around Davao City.

156 Chapter 5 Platystictidae of the Philippines

35 36

37

Figures 35-37. Drepanosticta furcata sp. n., male [JvT 18862, Siquijor, Barrio Bandilaan, 28-23.iv.1993]. – 35, pronotum, oblique view. – 36, anal appendages, dorsal view. – 37, idem, left lateral view. Scale bar 1 mm.

Drepanosticta furcata van Tol, sp. n. head bronze-black, especially on dorsum of head (Figs 35-37, 102) with coppery shine; transverse occipital carina well developed with sharp lateral extremities; antennae Drepanosticta sp. n. 4 (cf. lymetta). – Hämäläinen & Müller 1997: 257, 276 [Siquijor, 7 males in collection]. yellowish, flagellum brown (right flagellum missing in holotype). Type material. – Holotype male [JvT 18860] in RMNH: Thorax. Pronotum (Fig. 35) with anterior lobe pale ‘Philippines. Siquijor Id / Barrio Bandilaan / April 18.-23. brown, darker in centre, well-developed, with anterior 1993. / Th. Borromeo legit / Coll. Roland A. Müller’. – margin erect; middle lobe flat, yellowish brown; lateral Paratypes. Same data as holotype, 6 males (in RMNH). lobe brown as lower part of synthorax; posterior lobe brown, narrow, hind margin with paired process Diagnosis. – Medium-sized species with brown approximately twice as long as median length of pronotum; pronotum with a paired and forked process; posterior lobe, more or less erect or lying flat over distinguishable from other species in the D. lymetta synthorax, ending in a bifurcation of which inner group by the long inner branch of the forked process; branch is distinctly longer than outer brach (virtually the superior appendage with a subbasal, very sharp, symmetrical bifurcation in one specimen). Synthorax medio-ventrad tooth, a second and more distal tooth castaneous. Legs dirty yellow, only somewhat darker as present in D. lymetta is only discernable as a knob- at joints of femora (right fore leg missing in holotype). like structure; confined to Siquijor Id. See also remarks Wings hyaline, Px 14 in fore wing (left wing with under D. lymetta. two additional cross-veins), Px 14 in hind wing; origin of R4+5 at subnodus, IR3 halfway first cell Male [holotype, JvT 18860]. – Head. Labium brown; distal to that level; Arculus arising distinctly distal to labrum yellowish white, but anterior two-fifths pale Ax2; quadrangle of fore wing somewhat widening brown; mandibles yellowish white, with anterior posteriorly; anal veins shortly stalked, slightly one-sixth brown; anteclypeus bluish white, rest of asymmetrical; pterostigma castaneous, c. 2.4 times as

157 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

39 38 40

Figures 38-40. Drepanosticta hermes sp. n., male [JvT 18824, Mindanao, Davao Oriental, Boston, Mt Agtuuganon, 1020 m, 29.v-7.vi.1996]. – 38, pronotum, oblique view. – 39, anal appendages, dorsal view. – 40, idem, left lateral view. Scale bar 1 mm.

long as high; proximally obtusely angulate, distal side Drepanosticta hermes van Tol, sp. n. somewhat convex; cells between Costa and R1 distal to (Figs 38-40, 102) pterostigma rather frequently divided. Drepanosticta sp. n. 16. – Hämäläinen & Müller 1997: 258, Abdomen. Brown, but with paler yellowish white 277 (distribution Mindanao, Davao Oriental). markings as follows: ventro-anterior part of segment 2, anterior one-fifth of segment 3, anterior one-fourth Type material. – Holotype male [JvT 18850] in RMNH: of segment 4-6 (segment 6 somewhat darker yellow), ‘Philippines, Mindanao Id / Davao Oriental, Boston / Mt segment 10 pale brown. Appendages (Figs 36-37) Agtuuganon, Camp 55 / May 29. - Juni 7, 1996, 1020 m / Müller/Buenafe/Gorostiza leg. / Coll. Roland A. Müller’. yellowish brown; superiors in dorsal view stout in – Paratypes (43 specimens, all in RMNH): same data als basal half, on innerside ending in sharp tooth directed holotype, 42 males ; same, Boston, Caatijan, 500-550 m, ventro-axiad; distal half of superiors scope-like, 28.v.1996 (Müller, Buenafe & Gorostiza), 1 male. ending subacute; inferiors in ventral view with stout base, distal half straight, cylindrical, the tip strongly Diagnosis. – Relatively small, dark, castaneous constricted and outer margin curved 80˚ inward. brown species, closely allied to D. lymetta Cowley; Measurements. Hind wing 21.5 (21-22) mm, the processes of the hind lobe of the prothorax long, abdomen including appendages 33 (31-33) mm; one bifurcate at top with a short inner branch and a longer theratological specimen with abdomen of 24 mm. outer branch; the very slender superior appendages (dorsal view) diagnostic; base of superiors in lateral Female. – Unknown. view distinctly directed dorsad; ventral teeth of superiors with basalmost one approximately in middle Etymology. – Furcatus (Latin), forked; for the shape of of appendage, the other tooth poorly developed. See the processes of the hind margin of the posterior lobe also under D. lymetta Cowley. of the pronotum. An adjective. Distribution (Fig. 102). – Philippines Islands (West Male [JvT 18819]. – Head. Labium brown; labrum, Visayan region): Siquijor Island. mandibles and anteclypeus creamish white; anterior one-third of labrum pale brown, mandibles with

158 Chapter 5 Platystictidae of the Philippines

narrow anterior dark line; rest of head brownish black; will remain uncertain until pairs are found in copula. parorbital carina and postocular carina well developed Hämäläinen & Müller (1997: 277) mention 45 males. with distinct, acutely angulate lateral extremities; One specimen (JvT 18825) was identified as the antennae dirty yellow, flagellum brown. present species, but is in my view not conspecific, and Thorax. Pronotum (Fig. 38) laterally and medially concerns Drepanosticta flavomaculata sp. n. brownish black; rest dirty yellow to pale brown; Etymology. – Hermes (Greek), herald of the Olympian anterior lobe short, with anterior margin distinctly gods, usually depicted with a winged hat; for the erect; posterior lobe with a paired process on hind structure of the paired processes of the posterior lobe of margin, process as long as middle lobe, lying more or the pronotum. A noun in apposition. less flat over synthorax; bifurcate at top, with a short Distribution (Fig. 102). – Philippine Islands: tooth directed mediad, and a long tooth directed Mindanao (Davao Oriental). abaxiad, this tooth approximately as long as base of fork. Synthorax castaneous; some parts slightly paler, but no significant creamish white stripes; legs dark Drepanosticta lymetta Cowley yellow. Wings hyaline, venation brown; Px 17 in fore (Figs 41-43, 102) wing and Px 15 in hind wing; R4+5 arising from subnodus; IR3 halfway first cell distal to that level; Drepanosticta lymetta Cowley, 1936: 161-163, figs 13-14 Arculus just distal to Ax2; quadrangle widening distally (original description, holotype male, Philippine Islands, in forewing; anal veins subequal, stalked; pterostigma Surigao, Mindanao, C. F. Baker [BMNH] [examined]). – Needham & Gyger 1939: 263-264, figs 197-198, 200, width 1.5 times the height; several cells between Costa 202, 205, 310 (description, material from type locality); and R1 beyond pterostigma divided. Hämäläinen & Müller 1997: 257 (distribution Mindanao Abdomen. Brown, but anterior one-sixth to one-fifth subregion: Mindanao). of segment 3-7 somewhat paler; appendages (Figs 39-40) dark yellow; superior appendages slender, Material examined (all Philippines, in coll. RMNH, except base short, distal part forcipate with very short dorsal given otherwise). – Mindanao. Surigao, 21.v.1915 (G. tooth, and short and sharp ventral tooth in middle Boettcher) 1 female (ex SMFD); Surigao, 12.xi.1915 (G. Boettcher) 1 female in SMFD; Surigao, 23.viii.1916 (G. and a broader process distal to the sharp tooth, top of Boettcher) 1 male (ex SMFD) and 2 males 2 females in appendage blunt; inferiors with slender base, distal half SMFD; Surigao, before 1920 (C.F. Baker) 1 male (in BMNH) of inferior subcylindrical, somewhat bent outward, the [holotype of Drepanosticta lymetta Cowley]; Agusan [del Sur], top slender and directed inward, the tip curved dorsad. San Fran[cisco] (Quate) 1 female; Davao del Norte, Maco, Measurements. Hind wing 21.5 (21-23) mm, abdomen Mainit, Mait Creek, 500-700 m, 14-17.ix.1995 (A. Buenafe) 6 males; Davao del Norte, Maco, Mainit, Masara Mine, 500- including anal appendages 34 (33-34) mm. 700 m, 27-28.xi.1995 (A. Buenafe) 1 male.

Female. – Unknown (but see below under remarks). Diagnosis. – A rather inconspicuous brown species, the males characterized (but see below under remarks) Remark. – The female of this species is almost by a pair of shortly branched processes on the hind certainly among the specimens collected by Müller margin of the hind lobe of the pronotum, and rather and collaborators in Davao Oriental. However, straight superior anal appendages with halfway a dorsal Drepanosticta taurus, a species with similar coloration as tooth and two medially directed teeth at innerside, the present species, is syntopic. Females (8 specimens) of which the basalmost one is sharp. This species is collected on Mt Agtuuganon with coloration as D. closely related to D. hermes sp. n. from northern Davao hermes sp. n. or D. taurus Needham & Gyger have Oriental (Boston region), which differs by the much a very different hind margin of the hind lobe of the longer outer branch of the pronotal process, a more pronotum. Consequently, the status of these females slender base of the superior appendage (dorsal view),

159 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

41 43 42

Figures 41-43. Drepanosticta lymetta Cowley, male [JvT 18613, Mindanao, Davao del Norte, Maco, Mainit, 500-700 m, 14-17. ix.1995]. – 41, pronotum, oblique view. – 42, anal appendages, dorsal view. – 43, idem, left lateral view. Scale bar 1 mm. and by the inferior appendages, which are curved c. 1.8 times as wide as high, rhomboid; cells between rather than straight in the distal half. It is also close Costa and R1 posterior to pterostigma undivided. to D. furcata sp. n. from Siquijor, which differs by Abdomen. Brown, anterior one-tenth of segment a significantly longer inner branch of the pronotal 3-6 yellow, dorsally bluish white; appendages (Figs process, also by the absence of the dorsal tooth on the 42-43) yellow; superiors in dorsal view very stout, superior appendages, and a poorly developed second converging into a short, blunt dorsal tooth at circa ventral tooth on superior appendage. three-fifths from base, distal two-fifths laterally compressed, semicircular; at base of club-shaped top a Male. – Head. Labrum, anteclypeus, mandibles ivory pair of characteristic ventral teeth, the basal one sharp; white; anterior one-fourth of labrum brown; rest of inferiors in ventral view basal one-third rather stout, head bronze-black; transverse occipital carina distinct, distal two-thirds straight, subterminally constricted with distinctly angulate lateral extremities. Antenna at innerside, the tip curved nearly 90˚ mediad, hardly with scapus brown; pedicellus and flagellum yellowish directed dorsad. brown. Measurements. Abdomen incl. appendages 30-33 mm, Thorax. Pronotum (Fig. 41) predominantly brown, hind wing 20-21 mm. but sides of anterior and median lobe white; central part of posterior lobe and lateral lobes brownish black; Female. – As the male, but hind margin of posterior anterior, median and lateral lobes simple; posterior lobe of pronotum without projections. lobe with lateral portions narrow, with elongate Measurements. Abdomen 27 mm, hind wing 19 mm. stout process on posterior border, process ending in a terminal tooth at outer surface, approximately as Status. – The holotype ofDrepanostigma lymetta long as width of process; synthorax bronze-black, a Cowley is incomplete (abdominal segments 6-10 paler marking over humeral suture; legs yellow. Wings missing), while the only paratype, also a male, lacks hyaline, venation middle brown, but Costa somewhat head and abdominal segments 3-10. The incomplete darker; Px 15 in fore wing, 13 in hind wing; R4+5 status of both types was already so when this species arising from subnodus, IR3 arising half a cell distal to was described, so that the structure of the abdominal that level, Arculus distinctly distal to Ax2, anal veins segments and the anal appendages has remained shortly stalked, distinctly asymmetrical; pterostigma unknown. Material available for comparison was

160 Chapter 5 Platystictidae of the Philippines

collected in Surigao by G. Boettcher in 1915/1916, male diagnostic; superior appendages of male in dorsal but unfortunately, Drepanosticta material from view with stout base, the distal half slender. this area is not well represented in the R.A. Müller collection. The structure of the pronotum of a male Male. – Head. Labrum, mandibles and anteclypeus collected by Boettcher, now in the RMNH collection white or bluish white, anterior fourth of labrum with [JvT 19767], is rather close to the holotype (see also brown line; rest of head bronze-black, frons and vertex Cowley 1936, Fig. 13) and was identified as D. lymetta coriaceous; antenna with scapus brownish black, by Lieftinck. The structure of the anal appendages of pedicellus dirty yellow, flagellum brown; transverse the specimens from Davao del Norte: Maco agrees occipital carina distinct, with sharp triangular rather well with JvT 19767, but the structure of the extremities. rather characteristic processes of the hind lobe of the Thorax. Pronotum (Fig. 44) with anterior and median pronotum is distinctly different. The divided processes lobe, and lateral portions of posterior lobe brown, are better developed axiad than abaxiad in the type, rest brownish black. Anterior lobe large, erect at 45º; but in the specimens from Maco (e.g. JvT 18613) the lateral parts of median lobe somewhat protuberant; outer branch is distinctly larger than the inner branch. posterior lobe short, with lateral portions with very Also, the appendages of JvT 19767 are brown and sharp and long processes, directed dorsad, length of concolorous with the abdomen, while the appendages process of same length as median lobe of pronotum, of all Davao specimens are dirty white, and clearly both processes running more or less parellel. Synthorax distinct from the abdomen, including segments 9-10. castaneous, dorsally bronze-black; legs yellow. Wings The differences may prove sufficient to distinguish hyaline, venation brown; Px 17 in fore wing, Px 15 more than one taxon in due time. For the present, in hind wing; R4+5 at subnodus, IR3 approximately I have refrained from naming these forms, awaiting halfway first cell distal to that level; Arculus distal to further material from Surigao, and also from the region Ax2; quadrangle fore wing distinctly widening; anal between San Francisco (Agusan) and Maco. veins subequal, unstalked; pterostigma brown, with Distribution (Fig. 102). – Philippine Islands: narrow pale margin against veins, oblong, c. 1.7 times Mindanao region (Eastern Mindanao). as wide as high, the proximal side acutely angulate. Abdomen. Segments 1-6 brown, segments 7-10 brownish black, anterior portions of segments Drepanosticta taurus Needham & Gyger indistinctly paler; appendages (Figs 45-46) dark yellow, (Figs 44-46, 102) base of superiors dark grey, somewhat conical, distal Drepanosticta taurus Needham & Gyger, 1941: 145-146, half semi-circular, bent inwards, tip sub-squarish; at figs 1, 2, 5-7, 10 (original description, holotype male, base of basal portion three teeth as follows: one dorsal type locality Mindanao, Davao Province, La Lun river (in MCZC). directed caudad, a larger, sharp tooth directed inward Drepanosticta taurus Needham & Gyger. – Hämäläinen & just basal to first, and a broad and inconspicuous tooth Müller 1997: 258 [distribution Mindanao subregion: just distal to first; inferiors with stout base, distal part Mindanao). with base straight, just directed outwards, the top bent Drepanosticta sp. n. 17. – Hämäläinen & Müller 1997: 258, 90º inward, innerside emarginate, tip not directed 277 [distribution Mindanao]. dorsad. Measurements. Hind wing 21-24 mm, abdomen Material examined. – Philippines, Mindanao Id., Davao Oriental, Boston, Mt. Agtuuganon, Camp 55, 1020 m, 29.v. - including appendages 33-36 mm. 7.vi.1996, Muller, Buenafe & Gorostiza, 42 males 1 female. Female. – Females of this species and D. hermes sp. n. Diagnosis. – Medium-sized dark species; long and erect are indistinguishable. posterior projections of posterior lobe pronotum in the

161 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

44

45 46

Figures 44-46. Drepanosticta taurus Needham & Gyger, male [JvT 18765, Mindanao, Davao Oriental, Boston, Mt. Agtuuganon, 1020 m, 29.v-7.vi.1996]. – 44, pronotum, oblique view. – 45, anal appendages, dorsal view. – 46, idem, left lateral view. Scale bar 1mm.

Status. – The specimens here assigned toD. taurus specimens representing D. taurus. Needham & Gyger, do not fully agree with the original Distribution (Fig. 102). – Philippine Islands: description, and were identified as a new species by Mindanao (Davao Oriental). The type locality of D. Hämäläinen & Müller (1997). Although the present taurus, La Lun river, was not found in any atlas or specimens have the same characteristic projections gazetteer, and is not represented on the map. of the posterior lobe of the pronotum, the size of the specimens is very different from the type. Needham & Drepanosticta megametta group Gyger explicitly mention the very elongate abdomen, measuring 47 mm, and a hind wing 25 mm for the Transverse occipital carina distinct, with acutely male. However, I presume that the size of the abdomen angulate lateral extremities; anterior lobe of pronotum is a printing error for 37 mm, a size that would better erect, at least in D. centrosaurus sp. n. partly folded; fit the given measurements of wings, female abdomen, posterior lobe of pronotum with a pair of processes on and the measurements of the specimens in the Müller hind margin, very wide and broadly connected with collection. Unfortunately, I have not been able to study the posterior margin of the posterior lobe; pterostigma the type to confirm my supposition. According to distinctly wider than high; superior appendages slender Needham & Gyger, the type was deposited in MCZN. without dorsal tooth, but a sharp tooth directed Dr. Philip D. Perkins of the Museum of Comparative medially on innerside; inferior appendages slender and Zoology informs me (e-mail 28 May 2004) that the straight, the tip emarginate, curved 90˚ mediad. Based type is not present in MCZN collection. On his on the structure of the anal appendages, presumably suggestion, I also contacted Dr. James K. Liebherr the sister-group of the D. lymetta group. at Cornell University, where Needham worked, who Included species: D. centrosaurus sp. n. and D. informed me that the type could not be traced in that megametta Cowley. collection as well. Awaiting the recovery of the type, Distribution: Mindanao. and possible more material from the Davao region in Mindanao, I have decided to consider the present

162 Chapter 5 Platystictidae of the Philippines

Drepanosticta centrosaurus van Tol, sp. n. anteclypeus ivory white, anterior one-third of labrum (Figs 47-49, 103) pale brown, rest of head bronze-black, frons and vertex coriaceous; transverse occipital carina distinct, Drepanosticta sp. n. (14). – Hämäläinen & Müller 1997: 258, lateral extremities apiculate; antenna with scapus and 277 (distribution Mindanao, Surigao del Sur). flagellum brown, pedicellus dirty yellow. Thorax. Pronotum (Fig. 47) with central part of Type material. – Holotype male [JvT 18715] in RMNH: anterior, median and posterior lobe, and complete ‘Philippines, Mindanao Id / Surigao del Sur, Tago / Meme lateral lobe dark brown; lateral portions of anterior River, 100-300 m / June 12.-18., 1996 / Buenafe A. / and median lobe bluish white, lateral portions of Gorostiza A. leg / Coll. Roland A. Müller’. – Paratypes (all posterior lobe middle brown; anterior lobe large and Philippines, in RMNH, 98 males 19 females), Mindanao. Surigao del Sur: San Miguel, Suba River, 150-250 m, erect; median and lateral lobes simple; posterior lobe 13.iv.1995 (Müller, Buenafe & Gorostiza) 29 males 3 females; with lateral portions large, each approximately as large Tandag, Hitaub Creek, 500-600 m, 16-19.iv.1995 (Buenafe as half of median lobe, base ca. one-third of hind & Gorostiza) 26 males 2 females; Carmen, 2. Equipment margin of posterior lobe, distal half somewhat wider on Shop, km 9 and 11 Lanang Line, 500 and 600-650 m, 21- innerside, and significantly broadened on outer surface. 24.iv.1995 (Müller, Buenafe & Gorostiza) 1 male 2 females; Lingig, Mandus, 100 m, 27.v.1996 (Müller, Buenafe & Synthorax concolorous castaneous, without bronze Gorostiza) 3 males 1 female; same locality as holotype, 38 shine, dorsal part finely coriaceous. Legs yellow. Wings males 11 females; San Miguel, foot of Diuata Mountains, hyaline, venation brown; Px 15 in fore wing, Px 14 Castillo, Kagda-o Creek, 300-500m, 2-5.iv.1998 (Piamonte) in hind wing; R4+5 arising distal to subnodus, IR3 in 1 male. distal half of first cell distal to subnodus; quadrangle of fore wing slightly widening posteriorly; Arculus arising Diagnosis. – Small and dark species, characterized distal to Ax2; anal veins somewhat asymmetrical, by very broad processes of the posterior lobe of sessile; pterostigma brown, with paler margin against pronotum, the bases wider than the distance between veins, c. 1.6 times wider than high; one cell between the processes; superior appendages with a large laterally, Costa and R1 beyond pterostigma in left hind wing but hardly ventrally directed tooth at base of distal, divided, remaining cells undivided (divided cells also clasper-like part. rare in paratypes). Abdomen. Castaneous brown, base of segments 3-6 Male [JvT 18715]. – Head. Labrum, mandibles, dorsally with oblong ivory white spot. Appendages

47

49

48

Figures 47-49. Drepanosticta centrosaurus sp. n., male [JvT 18625, Mindanao, Surigao del Sur, Tandag, Hitaub Creek, 500-600 m, 16-19.iv.1995]. – 47, pronotum, oblique view. – 48, anal appendages, dorsal view. – 49, idem, left lateral view. Scale bar 1 mm.

163 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

(Figs 48-49) pale brown or yellow; superior appendages wide posterior processes of the posterior lob of the very slender, in dorsal view base more or less straight, pronotum; differs from D. centrosaurus sp. n. by the a conspicuous tooth on innerside halfway of superiors shape of the processes, which are subrectangular in D. (well visible in lateral view in the holotype, but megametta and distinctly laterally projected at the top inconspicuous in many other specimens), superiors in in D. centrosaurus; anal appendages of males of both distal half bent semicircular, ending abruptly squarish; species very similar. inferiors in ventral view with basal two-fifths stout, distal part narrow at base, then slightly widening, the Male [based on description in Cowley 1936]. – Head. distalmost part abruptly constricted, bent 60˚ inward, Labium pale brown, labrum and mandibles pale the tip acute. yellow with narrow anterior brown border; anteclypeus Measurements. Hind wing 20 (19-21) mm, abdomen pale yellow; rest of head black, with small exterior including appendages 31 (30-32) mm. brown spot against lateral ocelli; transverse occipital carina well developed, with acutely angulate lateral Female. – As the male, but posterior margin of extremities. Antenna with scapus and pedicellus posterior lobe of pronotum rather simple, with a yellowish brown, flagellum darker apically. paired, very short process, in shape more as a denticle Thorax. Pronotum (Fig. 50) brown, median lobe paler; with a group of setae, rather close to the middle of the anterior lobe erect, posterior lobe produced posteriorly lobe, hind margin laterally of processes emarginate, as two caudally directed processes, extending over then rounded towards lateral corner. synthorax, slightly convex, apical corners rounded, Measurements. Abdomen 26-28 mm, hind wing 18-20 only external corner slightly projecting; no specialised mm. tufts of setae on processes. Synthorax dark brown, mesepisternum with black stripe against dorsal carina, Etymology. – Centrosaurus (Latin), a genus of mesepimeron paler dorsally and anteriorly, extending dinosaurs, for the similary of the ‘collar’. A noun in on to the dorso-posterior corner of mesepisternum; apposition. metepisternum with pale yellow ovoid dorso-anterior Distribution (Fig. 103). – Philippines: Mindanao spot; about dorsal third of metepimeron pale yellow. (Surigao del Sur). Legs pale yellow. Wings hyaline, venation dark brown. Px 15 in fore wing, Px 13 in hind wing; R4+5 arising at subnodus, extending to beyond level of pterostigma; Drepanosticta megametta Cowley IR3 arising half a cell distal to subnodus; quadrilateral (Figs 50-51, 103) long, very slightly widened distally in fore wing, Drepanosticta megametta. – Cowley 1936: 163-167, figs 15- scarcely so in hind wing; anal veins forming a Y-shaped 22 (original description, type locality Philippine Islands, brace with very short stem; pterostigma brown, Surigao, Mindanao (BMNH)). Drepanosticta megametta Cowley. – Lieftinck, 1937: 72-74 subquadrangular, surmounting one cell. (D. ephippiata from Celebes considered most closely Abdomen brown; segments 1, 2, 8-10 without any to D. megametta); Needham & Gyger, 1939: 263, 264 pale markings, 3 to 7 with a complete basal yellow (key, no new material); Hämäläinen & Müller 1997: 257 ring extending further apicad dorsally than laterally, (distribution Mindanao subregion: Mindanao). occupying the basal 0.08 of segment 3, 0.10 of segment 4, 0.12 of segment 5, 0.13 of segment 6 and Material examined. – Mindanao, Surigao, 23.viii.1916 (G. Boettcher) 2 males (juv) in SMFD; Mindanao, Surigao 0.24 of segment 7. Anal appendages (Fig. 51) brown, (Baker) 1 male (holotype) in BMNH. superior appendages about three times the length of segment 10, inferiors subequal to superiors; superiors Diagnosis. – A medium-sized species; distinguishable in dorsal view basally broad with short mesal spine from most other Philippine Drepanosticta by the directed meso-ventrad at 0.22 of length, concave

164 Chapter 5 Platystictidae of the Philippines

Drepanosticta: other species

The following species here assigned to Drepanosticta do not share a unique character or character set with other species in the Philippines. Some of the species definitely have Bornean affinities, others apparently are sister to one of the species groups defined here. A more detailed phylogenetic analysis is needed to reveal their 50 relationships. For the time, all these species are taken together in the rest-group. The species are treated in alphabetical order. Included species: D. aurita sp. n. (Mindoro), D. ceratophora Lieftinck (Palawan, Balabac), D. lestoides (Brauer) (Samar, Panaon Id., Dinagat, Mindanao), D. malleus sp. n. (Mindanao), D. mylitta Cowley (Luzon, Samar, Leyte, Homonhon, Panaon), D. myzouris sp. n. (Luzon), D. paruatia sp. n. (Palawan), D. pistor sp. n. 51 (Sibuyan, Negros, Panay, Luzon ?), D. quadricornu sp. n. (Busuanga, Palawan).

Figures 50-51. Drepanosticta megametta Cowley, male Drepanosticta aurita van Tol, sp. n. [holotype]. – 50, pronotum, dorsal view. – 51, anal (Figs 52-54, 107) appendages, left lateral view. Scale bar 1 mm.

Drepanosticta sp. n. 12. – Hämäläinen & Müller 1997: 257- 258, 277 (Mindoro). exteriorly, becoming narrower and concave interiorly, convex exteriorly, hollowed out mesally, apices turned Type material. – Holotype male [JvT 22190] in RMNH: meso-caudad; inferiors convex exteriorly, apices turned ‘Philippines, Mindoro Id / Mindoro Or. Prov., Calapan / mesad, interiorly straight with a preapical mesal Comonal, Mt Tarugin, um [= c.] 350 m / July 17./26. 1990 / Adrian Gorostiza legit / Coll. R.A. Müller’. – Paratypes (all projection, thereafter strongly concave; in lateral view in RMNH). Mindoro. As holotype, 14 males 3 females; Mt superiors directed dorso-caudad basally, then sharply Halcon, 1000-1500 m, 2-20.v.1994 (N. Mohagan) 8 males caudad and slightly ventrad; inferiors almost straight, 3 females. apices turned mesad and slightly dorsad; no specialised Other material examined (all in RMNH). – Lubang Island. tufts of setae. Mindoro Occidental, Looc, Mt. Gonting, Sitio Gonting, 300- 500 m, 8-15.vii.1997 (C. Nazareno) 5 males 1 female. Female. – As the male, but the posterior projections of the posterior lobe of the pronotum shorter, Diagnosis. – Transverse occipital carina poorly subquadrangular, the top more rounded. See further developed, lateral extremities hardly discernable; Cowley (1936: 165-166). processes on anterior lobe of pronotum indistinct in specimens from Mindoro, with a distinct paired Note. – This species is not represented in the R.A. anterior projection in specimens from Lubang Island; Müller collection in RMNH. posterior lobe of pronotum with swollen lateral Distribution (Fig. 103). – Philippine Islands: portions of posterior lobe with long and sturdy setae; Mindanao (Surigao province). synthorax brownish black, without pale markings;

165 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

54

53 52

Figures 52-54. Drepanosticta aurita sp. n., male [JvT 18749, Lubang Island, Mt. Gonting, Sitio Gonting, 300-500 m, 8-15. vii.1997; not a paratype]. – 52, pronotum, oblique view. – 53, anal appendages, dorsal view. – 54, idem, left lateral view. Scale bar 1 mm. pterostigma somewhat wider than high; superior hind wing; Arculus distinctly distal to Ax2; anal veins appendages without teeth or protuberances; inferior more or less symmetrical, shortly stalked; pterostigma appendages in ventral view straight, only the very tip brown, subquadrangular, width 1.4 times the height; curved axiad (not dorsad). proximal corner angulate, the distal side somewhat convex. Male [JvT 22190, holotype]. – Head. Labrum and Abdomen. – Brownish black, basal one-eighth of anteclypeus bluish white, anterior border of labrum segment 3, one-sixth of segments 4-6, one-third of brown; mandibles pale brown, except for blue segment 7 yellowish; a basal spot at ventro-anterior innerbasal corner, rest of head bronze-black; transverse margin of segment 8; abdominal segments 8-9 occipital carina narrow with inconspicuous lateral distinctly broader than other segments, brownish extremities; antenna with scapus and flagellum brown, black. Appendages (cf. Figs 53-54) dirty yellow, base pedicellus dirty yellow. of inferiors greyish brown; superiors in dorsal view, the Thorax. Pronotum (cf. Fig. 52) with central part of top dorso-ventrally flattened, the scoop-like caudal half anterior, median and posterior lobe, and complete subrectangular; inferiors with stout base, distal half lateral lobe, brownish black; rest brown; anterior slender, top rounded, the tip acute, hollow. lobe simple, with anterior margin erect, but without Measurements. Hind wing 20 (20-22) mm, abdomen conspicuous projections (for specimens from Lubang, including appendages 33 (33-35) mm. see under variation); median and lateral lobes simple; posterior lobe with lateral third of both sides erect at c. Female. – As the male, but the posterior processes of the 60˚, stout, subrectangular, somewhat wider than high, posterior lobe of the pronotum somewhat shorter than hind margin densely covered with long and sturdy in the male, but also with long setae; last abdominal setae; synthorax fully bronze-black; legs yellowish. segments brownish black, with anal appendages Wings hyaline, venation brown, pale brown in basal yellowish white, valves long, surpassing the cerci. half; Px 16 in fore wing, Px 15 in hind wing; R4+5 Measurements. Hind wing 21-23 mm, abdomen 32-35 distal to subnodus in forewing, at subnodus in hind mm. wing; IR3 distinctly in distal half of first cell distal to subnodus in fore wing, a little beyond the middle in Variation. – Pterostigma is many males more squarish

166 Chapter 5 Platystictidae of the Philippines

than holotype, width c. 1.3 times the height. Males the tip curved medio-dorsad, with a subterminal from Lubang Island with anterior lobe of pronotum medially directed sharp tooth. (Fig. 52) medially with a broad, unpaired process, approximately as high as the median length of anterior Male. – Head. Labrum and anteclypeus ivory white, lobe, the upper margin of the process itself bifid, the anterior border of labrum brown, mandibles greyish corners laterally projected, acutely angulate. Females brown, rest of head bronze-black, narrow transverse with the projections on anterior lobe of pronotum less occipital carina with indistinct lateral extremities; distinct. antenna with scapus and pedicellus dirty white, Although specimens from Mindoro and Lubang flagellum pale brown. are distinct by the shape of the anterior lobe of the Thorax. Pronotum (Fig. 55) dirty yellow, with lateral pronotum, the Lubang form has not been named portion of median lobe, median portion of posterior separately, since no other structural characters seem to lobe, and lateral lobes brownish black; anterior lobe distinguish both populations. large, not distinctly erect, posterior lobe with median Etymology. – Auritus (Latin), eared; for the shape of portion diagnostic with long (unpaired) process, the processes of the posterior lobe of the pronotum. An smoothly curved dorsad, at least as long as median adjective. length of rest of pronotum, but with much variation Distribution (Fig. 107). – Philippine Islands: Mindoro, between specimens; lateral portion of posterior lobe Lubang Island. simple; synthorax castaneous, lower parts brownish black; legs yellow. Wings hyaline, venation brown; Px 14-15 in fore wing, Px 13-14 in hind wing; Drepanosticta ceratophora Lieftinck R4+5 arising at subnodus, IR3 arising half a cell (Figs 55-57, 107) distal to that level; Arculus arising just distal to Ax2, Drepanosticta ceratophora Lieftinck, 1974: 117-120, figs anal veins stalked in most specimens; pterostigma 3-5 (original description, holotype male Palawan, subquadrangular, proximal side 80º angulate, distal Mantalingajan, Pinigisan, 600 m (in ZMUC)). – Hämäläinen & Müller 1997: 257 (distribution Palawan). side slightly more convex; cells distal to pterostigma between Costa and R1 undivided. Abdomen. Segment 1 yellow, segment 2 brown, Material examined (all Philippine Islands, in RMNH). posteriorly somewhat paler, segments 3-6 brown, – Palawan. Mantalingajan, Pinigisan, 600 m, 1.ix.1961, each segment with anterior and posterior one-fifth Noona Dan Exp. 61-62, 1 female (incomplete, paratype) yellow, segment 7 brown with anterior third yellow, (ex ZMUC); Quezon, Lamakan, Magmuni Stream, 27.v. - 1.vi.1991 (Borromeo & Buenafe) 6 males 2 females; idem, segments 8-10 brownish black, dorsum of segment 10 Magmuni Stream, 1000 ft, 16-22.v.1992 (Borromeo T) 8 with irregular oval, blue marking. Appendages (Figs males 2 females; idem, Quezon, Malatgao, Bugon, ii.1994 (E. 56-57) yellow, slender; superiors with base far apart Vinciguerra & E. Horn) 1 female. – Balabac, Palawan Prov., and gradually tapering, distal three-fifths laterally Brgy. Dalahuan, 5-30 m, 8-14.ii.1997 (A. Buenafe) 7 males compressed, somewhat hollow; inferiors stout at base, 1 female. distal half slender, more or less straight, the tip curved Diagnosis. – Small species. Transverse occipital carina inward; subterminally a toothlike process on innerside, and lateral extremities indistinct; anterior lobe of directed 90˚ inward. pronotum simple without projections; posterior Measurements. Hind wing 17-18 mm, abdomen lobe with a single median spine, diagnostic within including anal appendages 28-30 mm. the Philippine Drepanosticta species; synthorax castaneous; pterostigma slightly wider than high; Female. – Coloration very similar to the male; posterior superior appendages long, slender and straight; inferior lobe of pronotum rounded without any indication of appendages also slender, the distal half hooked laterad, median posterior process; last abdominal segments

167 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

57

55 56

Figures 55-57. Drepanosticta ceratophora sp. n., male [JvT 20169, Balabac, Brgy Dalahuan, 5-30 m, 8-14.ii.1997]. – 55, pronotum, oblique view. – 56, anal appendages, dorsal view. – 57, idem, left lateral view. Scale bar 1 mm.

stoutly built, castaneous, but segment 10 and cerci Drepanosticta lestoides Brauer. – Needham & Gyger 1939: dirty white. 263, 264 (key, no new material); Lieftinck 1961: Measurements. Hind wing 17-18 mm; abdomen 25-26 133-135, fig. 47 (redescription, additional male from Surigao); Hämäläinen & Müller 1997: 257 (distribution mm. East Visayan region: Panaon, and Mindanao subregion: Mindanao, Dinagat). Affinities. – Although there is no other Philippine species of Drepanosticta known with an unpaired Material examined. – (All Philippine Islands, and in RMNH; erect posterior process of the pronotum, this species arranged per island, from north to south). – Samar, Samar Prov., Hinubangan [=Hinabangan], Arizona, 100-280 m, resembles the Bornean Drepanosticta monoceros 29.iii.-6.iv.1997 (R.A. Müller) 1 female, idem, San Isidro, Lieftinck, only known from one male and one female San Isidro river, 90-200 m, 31.iii.-5.iv.1997 (R.A. Müller) 1 specimen. The latter species, however, is very small, has female. – Panaon Id, San Francisco (various places), viii.1988 distinctly different coloration, while not only the male, (W. Catal) 3 males 2 females. – Dinagat Id., Surigao del but also the female is provided with a conspicuous Norte, Loreto, Mt Canbinlio, Canbinlio river, 4.vi.1988 median process on the posterior lobe of the pronotum. (A. Buenafe) 1 male. – Mindanao, Surigao, 30.x.1915 (G. Boettcher) 1 male; South Cotabato, Mt. Matutum, 500- Lieftinck considered D. monoceros a close relative of D. 700 m, 16-19.ix.1993 (Th. Borromeo) 1 male; Lanao del crenitis Lieftinck. Norte, Iligan, Tinago Falls. 8˚09’33”N 124˚11’11”E. 160 Distribution (Fig. 107). – Philippine Islands: Palawan, m, 25.iii.2004 (V. Kalkman & J. van Tol) 3 males; Davao Balabac. Oriental, Boston, Mt Agtuuganon, 1020 m, Camp 55, 29.v.- 7.vi.1996 (Müller, Buenafe & Gorostiza) 2 males 1 female; Mt Gantongan, Gantongan river, x.1991 (A. Buenafe) 1 Drepanosticta lestoides (Brauer) male; Surigao del Sur, Tabon, Tabon Falls, 100-200 m, 26.v., (Figs 58-60, 104) 8.vi.1996 (Gorostiza & Buenafe) 1 male.

Platysticta lestoides Brauer, 1868: 552-553 (original Diagnosis. – Large species with colourful thorax, description, type male, type locality Mindanao [in IRSN, pronotum with dirty white anterior, median and not examined]). – Selys 1882: 30 (no new records, dates provided). laterial lobes, and brownish black hind lobe, synthorax Platysticta (Platysticta) lestoides. – Selys 1886: 154-155 with wide yellowish white stripes over metepisternum (description). and metepimeron. Only other Philippine species of

168 Chapter 5 Platystictidae of the Philippines

59

58

60

Figures 58-60. Drepanosticta lestoides (Brauer), male [JvT 18620, Mindanao, Davao Oriental, Boston, Mt Agtuuganon, 1020 m, 29.v-7.vi.1996]. – 58, pronotum, oblique view. – 59, anal appendages, dorsal view. – 60, idem, left lateral view. Scale bar 1 mm.

Drepanosticta with similar coloration are D. moorei anterior portion of lateral lobe pale yellow, posterior Van Tol & Müller and D. luzonica sp. n. from Luzon; part of lateral lobe and most of posterior lobe dark appendages of both species, however, are distinctly brown, lateral portion of latter slightly paler; anterior, different, especially long tubercle directed mediad median and lateral lobes simple; posterior lobe short, halfway on inferior appendage of D. lestoides diagnostic flat, only lateral corners at hind margin with triangular (Figs 59-60). erect structure. Synthorax brownish black, but with Male. – Head. Labrum and anteclypeus bluish white, conspicuous bluish yellow transversal markings on but anterior one-sixth of labrum black, mandibles metepisternum against mesepimeron and lower half black except for bluish white inner basal corner; rest of metepimeron, apically continuing on lower hind of head bronze-black and very shining, microsculpture part of metakatepisternum. Legs yellowish, femora at only partly, very superficial and without conspicuous joints greyish brown. Wings hyaline, venation brown, longitudinal striae between eyes; hind margin of head but Costa brownish black in basal half and other main without transverse occipital carina, but parorbital veins pale brown in basal half; venation rather open; carina present; antenna with scapus brownish black, R4+5 arising at or just distal to subnodus, IR3 arising pedicellus and flagellum brown. just distal to that level; Arculus arising just distal to Thorax. Pronotum (Fig. 58) with anterior, median and Ax2; quadrangle only slightly widening in fore and

169 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

hind wing; anal veins asymmetrical and almost sessile; Drepanosticta malleus van Tol, sp. n. pterostigma brown with broad pale lines against (Figs 61-63, 107) veins, subquadrangular, width c. 1.5 times the height, Drepanosticta sp. n. 15. – Hämäläinen & Müller 1997: 258, proximal lower corner acutely angulate. 277 (Mindanao). Abdomen. Segments 1-7 pale brown, segments 1-2 ventrally yellowish, rest abdomen brownish black, Type material. – Holotype male [JvT 18794] in RMNH: ‘Philippines, Mindanao Id / Davao Oriental, Cateel, but basal half of segment 9 with a paired irregular Aliwagwag Falls, 200-300 m / April 10, 1995 / Buenafe / squarish blue marking. Appendages (Figs 59-60) Gorostiza legit / Coll. Roland A. Müller’. – Paratype: Same with superiors semicircular, the tip distinctly wider, data [head broken to pieces], 1 male (in RMNH). approximately halfway a dorsal tooth directed inward, its length approximately as diameter of appendage; Diagnosis. – Male (female unknown) unmistakable inferiors straight, the tips sharply bent inward and by its huge posterior processes of posterior lobe of touching, subbasally at ventral surface an inward pronotum; appendix inferior with subterminal inward directed triangular process, at dorsal surface a stout and directed tooth. The last character shared with D. long tubercle, directed inward and touching opposite myzouris sp. n., of which the structure of posterior tubercle. process is very different. Measurements. Hind wing 25-27 mm; abdomen including anal appendages 44-46 mm. Male [JvT 18794, holotype]. – Head. Labrum and anteclypeus ivory white, labrum with anterior third Female. – Generally similar in coloration to the male; light brown, mandibles greyish brown; rest of head structure of pronotum as male; abdominal segments brownish black, frons and vertex coriaceous; transverse with segment 1 yellowish white, dorsally a brown occipital carina distinct, with acutely angulate annule against hind margin connected with a paired triangular extremities; antenna with scapus brown, rest brown longitudinal line; segments 2-7 brown, the first of antenna dirty yellow. segments paler than the last, segment 2 with a narrow Thorax. Pronotum (Fig. 61) with anterior, median and longitudinal line over middle of segment and a broad middle part of posterior lobe middle brown, upperparts yellow stripe against sternite; segments 8-10 brownish somewhat darker; median portion of posterior lobe, black, segment 8 with a semi-circular spot against and lateral lobe brownish black; anterior lobe distinct sternite; valvifer brownish black, valve pale, dirty and erect, middle and lateral lobes simple, posterior yellow, reaching distinctly beyond cerci. lobe with lateral portions distinctly developed, nearly Measurements. Hind wing 25-27 mm, abdomen 37-40 as long as median line from anterior to posterior lobe, mm. base broad and long, the top abruptly connected to a transversal structure, strong and relatively short on Affinities. – Not closely related to any other Philippine innerside, longer and more slender outside, transversal Drepanosticta species. Within the Philippines, the structure with sturdy setae. Synthorax castaneous. Legs characteristic structure of the male appendages is yellow, femora against tibiae with a narrow brown ring. unique to the present species. D. lestoides (Brauer) Wings hyaline, venation brown; Px 16 in fore wing, seems to have closer affinities to species from Borneo, Px 15 in hindwing; R4+5 at subnodus, IR3 halfway e.g. D. actaeon Laidlaw, or the mainland of Southeast first cell distal to that level; Arculus just distal to Ax2; Asia, e.g. D. quadrata (Selys). quadrangle widening posteriorly in fore wing; anal Distribution (Fig. 104). – Samar, Panaon, Dinagat and veins distinctly stalked; pterostigma oblique, width Mindanao. A rather widespread but uncommon species c. 1.8 times the height, anterior and posterior side where found. parallel-sided, distal side somewhat convex; several cells between Costa and R1 distal to pterostigma divided.

170 Chapter 5 Platystictidae of the Philippines

61 62

63

Figures 61-63. Drepanosticta malleus sp. n., male [JvT 18793, Mindanao, Davao Oriental, Cateel, Aliwagwag Falls, 200-300 m, 10.iv.1995]. – 61, pronotum, oblique view. – 62, anal appendages, dorsal view. – 63, idem, left lateral view. Scale bar 1 mm.

Abdomen. Castaneous, basal one-tenth of segments Affinities. – The inward directed, subterminal tooth of 3-5 paler; anal appendages (Figs 62-63) long and the inferior appendage is shared, among the Philippine slender, superiors smoothly tapering from base to top, taxa, with D. myzouris sp. n. Clustering into one approximately halfway on innerside a sharp triangular group is unrealistic, since they do not share characters tooth, a shorter and broader protuberance just distal in the transverse occipital carina, the structure of to this tooth; tip of superiors rounded dorsally, acutely the pronotum and the pterostigma. The subterminal pointed ventrally; inferiors distinctly longer than tooth of the inferior appendages occurs widely in superiors, base stout, distal half nearly straight, the Platystictidae, including Drepanosticta, and has to be top 80˚ bent inward, slender; subterminally a short considered the plesiomorphic character state. triangular tooth directed axiad. Etymology. – Malleus (Latin), hammer; for the Measurements. Hind wing 21 mm, abdomen structure of the paired process of the posterior lobe of including appendages 32 (34) mm. the pronotum. A noun in apposition. Distribution (Fig. 107). – Philippine Islands: Female. – Unknown. Mindanao (Davao Oriental).

171 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

64 65 66

Figures 64-67. Drepanosticta mylitta Cowley, male. 64, pronotum, oblique view (‘typical D. mylitta’) [JvT 20141, Leyte, Mahaplag, Hilusig, Mt Balocaue, 700 m, 1.xii.1989]. – 65, pronotum, oblique view [JvT 21914]. – 66, anal 67 appendages, dorsal view [JvT 21914]. – 67, idem, left lateral view [JvT 21914]. Scale bar 1 mm.

Drepanosticta mylitta Cowley [in BMNH]; Las Navas, Brgy San Isidro, 100-350 m, 22- (Figs 64-67, 105) 28.v.1997 (A. Buenafe) 9 males 3 females. – Bilaran: Naval, Sayao Mtn, Horonan Creek, 200-400 m, 30.x.-3.xi.1992 (Th. Drepanosticta mylitta. – Cowley 1936: 160-161, figs 1-12 Borromeo) 2 males; Same, Sayao Mtn, Villaconsuelo, 4-7. (original description, holotype Philippine Islands, xi.1992 (Th. Borromeo) 1 male. –Leyte : Mahaplag, Hilusig, Borongan, Samar [BMNH] [examined]). – Needham & Balocaue Mtn, 600 m, 29.viii.-14.ix.1986 (Th. Borromeo) 7 Gyger, 1939: 263, 264 (key, no new material); Hämäläinen males 4 females; Same, 700 m, 29.xi.1989 (Th. Borromeo) 10 & Müller 1997: 257, 276 (distribution East Visayan males 2 females; Same, Magsuganao river, 18-30.ix.1990 (Th. subregion: Samar, Leyte, Biliran, Homonhon, Panaon; Borromeo) 20 males 5 females. – Homonhon: Magallanes Mindanao subregion: Dinagat; D. septima presumably a Point, 17.v.1988 (C. Treadaway) 1 male. – Dinagat: Loreto, synonym). Sinayao, June 1989 (A. Buenafe) 1 male. – Panaon: San ? Drepanosticta septima Needham & Gyger, 1939: 264-265, Francisco, Batong Lapad, viii.1988 (W. Catal) 2 males; same, figs 201, 204 (original description, holotype female, Samar, Kaneo Mtn, 400 ft (W. Catal) 1 male; Same, Gabing Gamay: 29 Apr 1924, R.C. McGregor [in CUIC?]). big river, x.1988 (W. Catal) 2 males 2 females; San Francisco, Drepanosticta sp. n. 6 (cf. mylitta). – Hämäläinen & Müller Anislagon Mtn, 100-150 m, 2.ii.1990 (Th. Borromeo) 1 1997: 257, 276. male; Same, Anislagon river, 10.x.1990 (Th. Borromeo) 3 males 2 females. Material examined (all Philippine Islands, all in RMNH, except specified otherwise). – Luzon, [Zambales], Subic Bay, Diagnosis. – Medium-sized, nearly completely 2.vii.1907 (J. C. Thompson) 1 male [ex CASC]; [Sorsogon], castaneous species. Pronotum typically with short and Bulusan, Mt Bulusan, San Francisco, Pagjasaan river, 400- 90º erect projections on hind margin of hind lobe, but 600 m, 4-5.xii.1997 (C.M. Nazareno) 2 males; Same, Obak some populations on Samar with these processes flat, river, 6.xii.1997 (C.M. Nazareno) 1 female; Same, Sioton river, 6-7.xii.1997 (C.M. Nazareno) 2 females; Same, San but otherwise structurally similar. Roque, Masacrot spring, 10-11.xii.1997 (C.M. Nazareno) 1 Male. – Head. Labrum and anteclypeus bluish white, male. – Samar, Borangan, date unknown, 1 male [holotype] anterior one-fourth of labrum brown; mandibles

172 Chapter 5 Platystictidae of the Philippines

greyish white, frons middle brown, a reniform middle- Female. – Similar to the male, but posterior lobe of brown spot posterior of lateral ocelli; rest of head pronotum with indistinct or even hardly discernable bronze-black, frons and vertex coriaceous; transverse spiny projection just before hind margin. occipital carina distinct with acutely angulate lateral Measurements. Hind wing 18-22 mm, abdomen 26-31 extremities; antenna with scapus and pedicellus very mm. pale brown, flagellum brown, the basal part somewhat paler. Variation. – A very variable species in the structure Thorax. Pronotum (Figs 64-65) with median parts of the posterior margin of the posterior lobe of the of anterior, median and posterior lobes dark brown, pronotum, and the anal appendages of the male. Males lateral parts creamish, lateral lobes dark brown; ‘typically’ have a short and paired spiny projection anterior, median and lateral lobes simple; posterior lobe just before the posterior margin of the posterior lobe, short, median portion with hind margin acute in the but some populations of Samar Id. (e.g. Hinabangan) middle, lateral portions with hind margin on innerside have the corners of the hind margin of the lobe slightly with an erect, slender, round process, approximately projecting, but are lacking the spine. These specimens as long as length of median axis of posterior lobe (see were considered a separate species by Hämäläinen & also Remarks, below). Synthorax castaneous, somewhat Müller (1997) (‘sp. n. 6’). The type of D. mylitta is obscured in lower frontal part. Legs yellowish white, also from Samar (Borongan, 11˚38’N 125˚27’E) and the coxae somewhat darker on posterior side. Wings is provided with spines on the hind margin of the hyaline, venation basal to nodus castaneous, rest dark posterior lobe. Presently, the variation of this taxon brown; Px 18 in fore wing, Px 16 in hind wing; origin within Samar is not understood. of R4+5 distal to or, more frequently, at subnodus; IR3 Also variation in the male appendages was observed. halfway first cell distal to that level; Arculus arising The width of the basal part of the superior appendages just distal to Ax2; quadrangle in fore wing somewhat (dorsal view) is very variable, but all populations widening posteriorly; anal veins sessile; pterostigma are characterised by the ventro-mediad tooth at the middle brown, subrectangular, width c. 1.8 time the base of the clasper-like part of the superiors; inferior height; cells between Costa and R1 undivided. appendages (ventral view) with distal part usually Abdomen. Middle brown, segments 3-7 with base of straight, only the tip distinctly tapering, hooked segments dorsally with a bluish white pale marking, mediad, the very tip dorsad; in some specimens, subtriangular (segment 3) or panduriform (spear- however, the distal halves diverging, but otherwise like) (other segments). Appendages (Figs 66-67) dark similar. In some populations (e.g. Leyte Id, Mt yellow to lightbrown, superiors in dorsal view with Balocaue) the male inferiors are very robust, viz. stout basal two-fifths, distal three-fifths much more terminal part c. two times as long as wide, as compared slender, curved outward, then inward, the tips not to c. four times in ‘typical’ specimens). quite touching, subsquarish, base of distal portion with These differences may indicate that D. mylitta a short inward directed, very sharp tooth; inferiors as actually consists of a group of closely related species. long as superiors, in ventral view with basal third stout, More material is needed to map the geographical distal two-thirds in basal half straight, rest forming a distributions of all character-states, and to re-evaluate heart-shaped figure with the other appendage, the top the value of the documented variation, while also bent 90˚ inwards, the tip curved dorsad. a molecular study of this species may contribute to Measurements. Hind wing 18-21 mm; abdomen understand the morphological variation. including appendages 28-32 mm; some specimens Distribution (Fig. 105). – Philippines Islands (from from Samar without spiny projections on hind margin north to south): Luzon, Samar, Biliran, Leyte, of posterior lobe of pronotum are larger, hind wing up Homonhon, Dinagat, Panaon. Widespread species, to 22 mm, abdomen up to 34 mm. including several small islands.

173 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

Drepanosticta myzouris Van Tol, sp. n. (Figs 68-70, 107)

Type material. – Holotype male [JvT 22124] in RMNH: ‘Philippines, Luzon Id / Camarines Sur, Pili / Bungcao Curry, 200-400 m / Mt Isarog, Caririca River / August 4.-15. 1997 / Celso Nazareno legit / Coll. Roland A. Müller’. – Paratypes (all in RMNH). Same data as holotype, 3 females.

Diagnosis. – Relatively large species, with distinct pale markings in posterior part of synthorax. Male unmistakable by its remarkable, sucker- or proboscis- like posterior processes of the posterior lobe of the pronotum; the same structure absent in female. Appendix inferior of male with a subterminal inward 68 tooth (for interpretation of phylogenetic relevance, see D. malleus sp. n.).

Male [JvT 22124, holotype]. Head. Labrum and anteclypeus pale blue, anterior border of labrum light brown; mandibles light brown; rest of head bronze- black, small blue marking at abaxial side of lateral ocelli; transverse occipital carina without extremities. Antenna with scapus pale brown, pedicellus dirty yellow with the tip castaneous, flagellum castaneous. Thorax. Pronotum (Fig. 68) with anterior, median, upper part of lateral and posterior lobes dirty yellowish white; lower part of lateral lobes brown; anterior, median and lateral lobes simple; innerside of lateral 69 portion of posterior lobe with a broad and flat process ending in a disc; length of process approximately median length of median lobe. Legs dirty white. Synthorax light brown, but an irregular brownish black stripe against dorsal carina, and bluish white, more or less rectangular markings in posterior quarter of mesepimeron, metisternum and metepimeron. Wings hyaline, venation rather open, brown; Px 13 in fore wing, Px 12 in hind wing; R4+5 arising at level of subnodus, IR3 arising just distal to that level; Arculus well distal to Ax2; quadrangle hardly widening

70 Figures 68-70. Drepanosticta myzouris sp. n., male [JvT 22124, holotype]. – 68, pronotum, oblique view. – 69, anal appendages, dorsal view. – 70, idem, left lateral view. Scale bar 1 mm.

174 Chapter 5 Platystictidae of the Philippines

posteriorly, anal veins symmetrical; pterostigma brown, Diagnosis. – Male unmistakable among the described with narrow paler lines against the veins, width in fore Drepanosticta of the Philippines. Large, with wing ca. 1.1 times the height, subtrapezoid, proximal conspicuous antehumeral stripe, remarkable pending side only somewhat shorter than distal side, in hind appendages of the posterior lobe of the pronotum, and wing with proximal side more oblique, shorter than the structure of the appendages with a clavate mediad distal side; a few cells between Costa and R1 divided in directed tubercle on innerside of superior appendage. one wing only. Not closely related to any of the other species of Abdomen. Segments 1-6 middle brown, basal one- Drepanosticta from the Philippines, but sharing several tenth tot one-eighth of segments 2-5 ivory white, characters with Bornean species, as Drepanosticta segments 8-9 castaneous, segment 10 dorsally blue, dupophila Lieftinck. Only one specimen known. ventral side of tergites brown. Appendages (Figs 69-70) brown, but dorsum of basal two-thirds of superiors Male [JvT 20337, holotype]. Head. Labrum, blue; base stout, the distal half laterally compressed, anteclypeus and inner basal corner of mandibles high, top subsquarish, a triangular tooth at base of bluish white; anterior two-fifths of labrum and larger club-shaped distal part; inferiors as long as superiors, part of mandibles black; rest of head bronze-black, in ventral view with stout base, the distal half smoothly microsculpture very superficial, dorsal part of head curved outward, than inward, the tip curved dorsad; very shining; transverse occipital carina distinct, with subterminally a long, sharp, inward directed tooth. lateral extremities apiculate. Antenna with scapus and Measurements. Hind wing 25 mm; abdomen including pedicellus dirty yellow, flagellum broken. appendages 38 mm. Thorax. Pronotum (Fig. 71) with anterior, median and lateral lobes ivory white, main part of posterior Female. – As the male, but process on posterior lobe lobe black, the lateral portion brown; anterior, of pronotum simple and relatively short, the top bent median and lateral lobes simple, posterior lobe short, inward, approximately the length of median line of the extreme lateral corners narrow based, elongate posterior lobe; abdominal segments with segments appendage directed ventrad, reaching beyond 8-9 brown, segment 10 dorsally pale blue, the anal ventral side of lateral lobe, and not immediately appendages ivory white; valves long, surpassing tip of conspicuous. Synthorax bronze-black with extensive appendages. pale markings as follows: wide antehumeral blue stripe Measurements. Hind wing 26 mm; abdomen 37 mm. anteriorly starting the width of stripe from anterior side of mesepisternum, length approximately three- Etymology. – Myzouris, sucker (Gr.), for the structure quarters mesepisternum; lower posterior corner of of the projection of the posterior process of the mesokatepisternum ivory white, as well as complete pronotum. A noun in apposition. metepisternum except for a triangular black marking Distribution (Fig. 107). – Philippine Islands, Luzon, in posterior corner, and complete metepimeron, Mt Isarog. Only known from the type locality. except for a narrow brown line against metapleural suture; legs yellow, brown rings around joints on femora and tibiae. Wings hyaline, venation brown up Drepanosticta paruatia van Tol, sp. n. to nodus, rest brownish black; Px 14 in fore wing, Px (Figs 71-73, 107) 13 in hind wing; R4+5 arising at or a trifle distal to Drepanosticta sp. n. 19. – Hämäläinen & Müller 1997: 258, subnodus, IR3 arising well distal to that level; Arculus 277. arising distinctly distal to Ax2, quadrangle somewhat

Type material. – Holotype male [JvT 20337] in RMNH: widening posteriorly in fore wing; anal veins somewhat ‘Philippines, Palawan Id / Port Barton, Waterfalls / May 28, asymmetrical in both fore and hind wing, shortly 1991 / Roland A. Müller legit / Coll. R. A. Müller’. stalked; pterostigma short, width 1.3 times height,

175 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

71

72

73

Figures 71-73. Drepanosticta paruatia sp. n., male [JvT 20337, holotype]. – 71, pronotum, oblique view. – 72, anal appendages, dorsal view. – 73, idem, left lateral view. Scale bar 1 mm.

all sides indistinctly convex, proximal sides acutely Female. – Unknown. angulate. Abdomen. Segment 1 ivory white, segments 2 to 7 Etymology. – Paruatia (latinized version of Greek), mostly brown, but segment 2 ivory white in lower part with hanging ears; for the structure of posterior lobe of of tergite, and narrow ivory white rings anteriorly on the pronotum. An adjective. segment 3-6; anterior three-quarters of segment 8, and Distribution (Fig. 107). – Philippine Islands: Palawan. anterior half of segment 9 pale, presumably blue in life, segment 10 completely brownish black, sternites of Drepanosticta pistor van Tol, sp. n. segments 8-9 blue. Appendages (Figs 72-73) brown, (Figs 74-76, 106) basal parts dark brown; both superior and inferior appendages long and slender; superiors with basal one- Drepanosticta sp. n. 13. – Hämäläinen & Müller 1997: 258, 277 [distribution Negros, Panay, Sibuyan] fifth relatively stout, ending in a bifurcation, consisting of an inward directed process approximately the Type material. – Holotype male [JvT 20260] in RMNH: length of basal part of superior; main stem of superiors ‘Philippines. Negros Id. / Negros Occ., Murcia / Mt Mawa, curving smoothly towards each other, the distal one- Brgy Canlandog / Balantak River, 400-500 m / May 4.-8. fourth widening ventrally; inferiors also with stout 1995 / Alex Buenafe legit / Coll. Roland A. Müller’. – base, distal three-quarters curved semi-circularly, the Paratypes (25 specimens, all Philippine Islands, in RMNH, arranged per island in chronological order). Sibuyan. top wider and scooplike, subterminally a very slender Romblon province, Magdiwang, Mt Guiting-Guiting, 300 tooth directed inward, a trifle longer than width of m, 22-23.vi.1986 (leg. ?) 1 male 1 female; Magdiwang, stem. Tampayan, Pawala river, 50-200 m, 19-31.vii.1986 (R.A. Measurements (type only). Abdomen including Müller) 2 males 1 female; Magdiwang, Silum, 5-100 m, appendages 39 mm; hind wing 21 mm. 3.viii.1986 (R.A. Müller) 1 female; Magdiwang, Tampayan,

176 Chapter 5 Platystictidae of the Philippines

7476 75

Figures 74-76. Drepanosticta pistor sp. n., male [JvT 18799, Negros Occ., Murcia, Mt. Maura, Balantak river, 4-8.v.1995]. – 74, pronotum, oblique view. – 75, anal appendages, dorsal view. – 76, idem, left lateral view. Scale bar 1 mm.

Camp Ga-ong, 80-150 m, 18-31.iii.1987 (R.A. Müller) 3 and vertex coriaceous; transverse occipital carina males 1 female; Magdiwang, foot of Mt Guiting Guiting, distinct, but without lateral extremities. Antenna with 200 m, 20-25.ii.1997 (ex coll. Treadaway) 1 male. – Negros. scapus and pedicellus dirty yellow, flagellum (damaged Negros Occidental Prov., Murcia, Mambucal, 2000 ft, 1-31. in holotype) castaneous. viii.1987 (A. Buenafe, C. Carzon) 2 females; Murcia, Mt Canlaon, Pula River, 3000 ft, 19.iv.1988 (A. Buenafe) 1 male; Thorax. Pronotum (Fig. 74) with following coloration: same, Asia River (no altitude given), 13.iv.1988 (A. Buenafe) anterior, median and posterior lobe dirty yellow, some 2 females; same, Pula River, ix.1990 (A. Buenafe) 1 male; parts more obscure; lateral lobes brown; anterior, Murcia, Mt Mawa, Brgy Canlandog, Balantak River, 400-500 median and lateral lobes simple; posterior lobe in m, 4-8.v.1995 (A. Buenafe) 1 male; same, Intalocan Creek, lateral corners with bifurcate appendage, anterior 800-900 m, 26.v.1995 (A. Buenafe) 1 male. – Panay. Antique province. San Remigio, Aningalam, Mt Clara, 600-850 m, portion narrow and directed abaxiad, posterior portion 28.vii-10.viii.1996 (A. Buenafe) 1 male 3 females; idem, Mt twice as long and wide, at base running abaxiad, Clara, 600-850 m, 28 Jul-10.viii.1996 (A. Buenafe) 2 females. distal part bent ventrad; synthorax bronze-black with Excluded from type series. – Luzon. Camarines Sur, Pili, dirty yellow parts as follows: dorsal carina, an oblong Bungcao Curry, Mt Isarog, Caririca River, 200-400 m, 4-15. triangular marking in distal three-quarters of both viii.1997 (C. Nazareno) 3 males 3 females; same, Himaao sides of humeral suture, metastigma, and another River, 200-400 m, 4-15.viii.1997 (C. Nazareno) 1 male 3 females. oblong triangular marking on posterior part of metepisternum. Legs yellow. Wings hyaline, venation Diagnosis. – Medium-sized, variegated species; differs brown; Px 13 (13-14) in fore wing, Px 13 in hind from other Philippine Drepanosticta by the simple wing; R4+5 at subnodus, IR3 halfway first cell distal structure of the anal appendages of the male without to that level; Arculus distinctly distal to Ax2; anal veins any teeth or tubercles, and the cleft sides of the symmetrical, not stalked (even divided in left fore wing posterior projections of the posterior lobe of pronotum of holotype); pterostigma brown, width 1.5 times the (see below for variation between populations). height; proximal side acutely angulate; first cell distal to pterostigma between Costa and R1 large; all cells in Male [JvT 20260, holotype]. – Labrum, mandibles that area undivided. and anteclypeus bluish white; narrow black line along Abdomen. Segments 1-2 and 8-10 dark brown, other anterior border of labrum; rest of head bronze-black segments middle brown, with narrow ivory-white with longitudinal striae between eyes, clypeus, frons markings in anterior one-tenth of segment 3-6, and

177 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

anterior two-fifths of segment 7. Appendages (Figs Borromeo) 2 males 1 female. – Palawan, Port Barton, 75-76) yellow, base of inferiors obscured; superiors 50-100 m, 15.vi.1985, [R.A. Müller], 1 female; Quezon flat, in dorsal view more or less parallel-sided over full district, Malatgao, Magmuni stream, 25-27.v.1991 (M. Hämäläinen) 1 male; Quezon, Lamaken [= Lamakan], length, distal two-fifths bent c. 60˚ towards each other; Magmuni stream, 27.v.-1.vi.1991 (Borromeo / Buenafe) 1 in lateral view distal part bent ventrad; inferiors with female; Port Barton, waterfalls, 28.v.1991 (R.A. Müller) 4 base stout, distal two-fifths narrow, base straight, the males 2 females; Mt Capoas, Sitio Caoban, Camp Neutcio, tip bent 80˚ towards each other. ca 280 m, 11-15.iii.1992 (Vinciguerra / Gorostiza) 2 females; Measurements. Hind wing 19 (18-21) mm; abdomen Mt Capoas, Bo Bandanan, Camp Inunugan River, 16-22. iii.1992 (Vinciguerra / Gorostiza) 1 male 4 females; Mt including appendages 32 (26-32) mm. Saint Paul, Tagabinit, Babuyan River, ca 160 m, 25-30. iii.1992 (Vinciguerra / Gorostiza) 2 males; Idem, ca. 350 m, Female. – Similar to the male, but more robust; 27.iii.1992, 1 male. structure of pronotum as in male; last abdominal segments stout, segment 9 subsquarish in lateral view; segments 8-10 brown; cerci dirty white; valves long, Diagnosis. – Small species, with distinct projections reaching at or beyond tip of cerci. on both the anterior and the posterior lobes of the Measurements. Hind wing 19-21 mm, abdomen 29-31 pronotum and pale stripe over metepisternum; other mm. species with two pairs of processes on the pronotum are D. belyshevi Hämäläinen, D. trachelocele sp. n., and Variation. – Specimens from Luzon are similar in most D. moorei Van Tol & Müller. Apart from structural respects, but some differ from the type in the structure differences in the projections of the pronotum, both D. of the posterior process of the posterior lobe of the belyshevi and D. trachelocele have distinctly clasperlike pronotum with the cleft dividing the lateral border of superior appendages, while D. moorei can most easily the process more equally. be distinguished on the pale stripe over the synthorax Females. – Females are remarkably well represented in that extends over metepisternum and metepimeron. D. the collection. quadricornu also differs from Philippine congenerics Etymology. – Pistor (Latin): miller; named after Roland by the structure of the anal appendages of the male; in A. Müller, St. Gallen; Müller is German for miller. A lateral view the superior appendage is strongly convex noun in apposition. with a subterminal, medio-ventrally directed triangular Distribution (Fig. 106). – Philippine Islands: Sibuyan, tooth. Negros, Panay and Luzon. A rather widespread species, but apparently uncommon where found. Male [JvT 20341]. – Head. Labrum, mandibles and anteclypeus ivory white, a narrow black anterior line on mandible; rest of head black and very shining Drepanosticta quadricornu van Tol, sp. n. without microsculpture; transverse occipital carina (Figs 77-80, 107) indistinct, without extremities. Antenna with scapus Drepanosticta sp. n. 20. – Hämäläinen & Müller 1997: 258, and pedicellus creamish white, flagellum castaneous. 277 (Palawan, Busuanga). Thorax. Pronotum (Figs 77-78) with anterior,

Type material. – Holotype male [JvT 20349] in RMNH: median and lateral lobes dark yellow, posterior lobe ‘Philippines, Palawan Id / Mt Capoas, Bo Banbanan / brown; anterior lobe erect, large, lateral corners Camp Inunugan River / 16./22. March 1992 / Vinceguerra/ with sharp process, approximately twice as long as Gorostiza leg.’. – Paratypes (all specimens Philippines, height of anterior lobe, directed dorsad; median and in RMNH) by island in chronological order. Busuanga lateral lobes simple, posterior lobe short, but lateral island, Busuanga, 1.viii.1990 (Th. Borromeo Sr) 1 male portions with long process, base directed caudad, (incomplete); Coron, 2.viii.1990 (Th. Borromeo Sr) 1 female; Coron, Mabentangen river, 4-7.v.1991 (Th. distal half curved upright, or just somewhat anteriad.

178 Chapter 5 Platystictidae of the Philippines

femora and joints of femora and tibiae. Wings hyaline, venation brown in basal part up to nodus, rest brownish black; Px 15 in fore wing, Px 13 in hind wing; R4+5 at level of subnodus, IR3 halfway first distal to that level; Arculus just distal to Ax2; quadrangle somewhat widening posteriorly in fore wing only; anal veins sessile; pterostigma brown, with narrow pale lines against the veins, oblong, width 1.8 77 times the height, proximal side acutely angulate; cells between Costa and R1 undivided (rarely divided in other specimens). Abdomen. Segment 1 ivory white; segment 2-8 with extensive ivory white rings as follows: anterior one- fifth of segment 2, anterior and posterior one-fifth of segments 3-6, segment 8 ivory-white in anterior two-fifths; segments 9 and 10 brownish black. Anal appendages (Figs 79-80) brownish black, convex; superiors in dorsal view with relatively slender base, gradually tapering towards acute tip, inner lateral 79 view shows tip as a long process on solid base, which 78 abruptly ends and bends ventrad; inferiors with stout base, rest nearly straight, the tips bent 50˚ twoards each other and nearly touching. Measurements. Hind wing 17.5 mm, abdomen including appendages 28 mm.

Female. – Very similar to male, including structure of pronotum, and coloration; all specimens in collection with abdomen discoloured, but presumably abdominal 80 segment 8 pale, segment 9 brown, segment 10 dark brown. Valves long and slender, surpassing level of cerci. Figures 77-80. Drepanosticta quadricornu sp. n., male [JvT Measurements. Hind wing 17-18 mm; abdomen ca 25 20343, Palawan, Port Barton, 28.v.1991]. – 77, pronotum, mm. oblique view. – 78, pronotum, dorsal view. – 79, anal appendages, dorsal view. 80, idem, left lateral view. Scale bar Affinities. – The sister-species of D. quadricornu sp. n. 1 mm. is presumably to be found on Borneo; further studies are needed. Synthorax castaneous, but posterior three-quarters Etymology. – Quadricornu (Latin), four horns; for of metepisternum with wide bluish-white stripe, the structure of the anterior and posterior lobes of the anteriorly sharp, posteriorly nearly touching hind pronotum. A noun in apposition. margin of metepisternum, ventrally a narrow brown Distribution (Fig. 107). – Philippines: Palawan and stripe along metapleural suture; posterior quarter Busuanga. of metepimeron white. Legs yellow, dark rings on

179 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

Protosticta Selys anteclypeus bluish white; narrow brownish black line

Protosticta Selys, 1885: cxlvi-cxlvii. along anterior border of labrum, approximately one- tenth of height of labrum, tapering towards corners The genus Protosticta Selys is traditionally distinguished and not quite reaching lateral corners; rest of head from other Platystictidae by combination of a straight, bronze-black, coriaceous; transverse occipital carina rather than fractured, IR3 vein, and the absence of well developed without lateral extremities; antenna lost. an Anal bridge (only the anal crossing is present). Thorax. Pronotum (Fig. 81) with anterior lobe brown Three platystictids with characteristic and similar in median one-third, lateral to that marking a small anal appendages, but differing in the vein venation, reddish brown triangular projection from anterior are assigned here to the new genus Sulcosticta. border of lobe directed posteriad, lateral quarters of Three other species with wing venation as the type lobe bluish white; median lobe with anterio-median species of Protosticta Selys, are described here as depressed part brownish black, the lateral portions Protosticta. They are apparently nor closely related to P. vaulted, brown on mediad surface, rest bluish simplicinervis Selys, nor to each other. As indicated in white, somewhat darkening posteriorly; lateral lobes the introduction, a more definitive arrangement has to brownish black, median portion somewhat paler; await a detailed phylogenetic analysis. posterior lobe medially with triangular brownish Included Philippine species: P. annulata (Selys), P. black marking, lateral thirds dirty yellow, with erect, lepteca sp. n., P. plicata sp. n. triangular, auriculate projections, approximately as Distribution: Luzon. long as median line of posterior lobe. Synthorax with dorsal carina anteriorly with a small yellow marking, mesepisternum black, a narrow dirty yellow line against Protosticta annulata (Selys) humeral suture and dorsal side of mesekatepisternum; (Figs 81-83, 108) mesokatepisternum black with a dirty yellow stripe Platysticta (Platysticta) annulata Selys, 1886: sep. 156-157 against coxa; mesepimeron black, a dirty yellow (original description, type female, type locality Luçon (in line against humeral suture, starting anteriorly the coll. Selys, IRSN). Platysticta annulata Selys. – Selys 1891: 218 (considered close distance of dorsal side of mesokatepisternum from relative of rufostigma from Borneo). posterio-dorsal corner of mesokatepisternum, running Protosticta annulata Selys. – Needham & Gyger 1939: 263, along suture, posteriorly covering posterior margin 264 (key, no new material). of mesepimeron, continuing anteriorly along first Drepanosticta annulata (Selys). – Lieftinck, 1961: 135-136 suture almost to level of metastigma; metepisternum (two females left, presumably not conspecific, no 577 with longitudinal, dirty yellow stripe from anterior- selected as lectotype, number 588 paralectotype; both in poor condition and status unclear); Hämäläinen & Müller ventral corner to metastigma, and another pale stripe 1997: 257, 276 (distribution Luzon). arising just posterior to metastigma, anteriorly acute, widening posteriorly, not quite reaching posterior side Material examined. – Luzon, Nueva Viscaya, Sta Fe, Dalton of synthorax; metepimeron black with rectangular Pass area, 900 m, 8-17.viii.1991 (A. Gorostiza) 1 male. bluish white marking against posterior margin. Legs dirty white, hardly visible darker rings on femora, Diagnosis. – Small and delicate species, synthorax with approximately one-quarter from joints with tibiae. longitudinal pale stripes as in taxa here assigned to Wings hyaline, venation brown, the posterior half Sulcosticta, last abdominal segments of male swollen; darker; Px 12 in fore wing, Px 12 in hind wing; origin posterior margin of posterior lobe of pronotum with a of R4+5 at subnodus, IR3 half a cell distal to that paired, triangular projection. level; Arculus arising well distal to Ax2; quadrangle somewhat widening posteriorly in fore wing; Ab vein Male [JvT 20075]. – Head. Labrum, mandibles, absent in all wings; pterostigma subquadrangular,

180 Chapter 5 Platystictidae of the Philippines

81 83 82

Figures 81-83. Protosticta annulata (Selys), male [JvT 20075, Luzon, Nueva Viscaya, Sta Fe, Dalton Pass area, 900 m, 8-17. viii.1991). – 81, pronotum, oblique view. – 82, anal appendages, dorsal view. – 83, idem, left lateral view. Scale bar 1 mm. proximal side oblique, width 1.3 times the height; cells to Selys when describing this taxon with ‘le rudiment between Costa and R1 distal to pterostigma undivided. du secteur inférieur du triangle manquant tout à fait, Abdomen brown, segments 1-2 castaneous, segments comme chez la Protosticta simplicinervis’ was no longer 3-6 light brown, segments 7-10 brownish black, available at all. with basal creamish white annulae as follows (lateral This information is valuable. One male platystictid view): segments 2-3 one-tenth of segment, segment in the Müller collection (data as above [JvT 20075]) 4 anterior one-eleventh, segment 5 one-fifteenth, was identified by M. Hämäläinen as the first male segments 6-7 one-twentieth, segment 8 half; segments of Drepanosticta annulata (Selys). However, the 8-9 much wider than other segments. Appendages (Figs wing venation of this male is clearly as described for 82-83) with creamish white superiors, and inferiors Protosticta, i.e. with missing anal bridge. This specimen with dark base and yellowish white terminal half; is here described as the male of D. annulata, but its superiors rather slender in dorsal view, a short tooth status can only be ascertained after males and females at c. 40% from base, distal part curved inward and from one site will be available, and can be compared ventrally widened; inferiors straight, smoothly tapering with the lectotype. towards the top, the top sharp and curved dorsad. Needham & Gyger already placed Platysticta annulata Measurements. Hind wing 17 mm, abdomen including Selys in the genus Protosticta, based on the note by appendages 27 mm. Selys on the state of the anal bridge. However, they clearly missed the first word of the sentence ‘... L’un Female. – No additional data. deux présente une anomalie singulière: le rudiment .... [as above]’, or: ‘One of them has an aberration ...’. Status of P. annulata. – Lieftinck (1961) re-examined Affinities. – Based on structure of the pronotum the type material in the Selys collection, and found that and the coloration of the synthorax, this species is the type series consisted of two very immature females. considered the sister-group of the species here included The poor status of preservation of these specimens in Sulcosticta gen. n. A further phylogenetic analysis made it impossible to match the selected lectotype with has to reveal whether the more simple structure of the any of the described or undescribed species known to inferior appendage of P. annulata is plesiomorphous Lieftinck. Besides, the two remaining specimens were or apomorphous compared with the structure in not conspecific. The second female, a paralectotype Sulcosticta gen. n. If it will appear to be an apomorphy, of D. annulata, is a female of Drepanosticta moorei P. annulata should be included in Sulcosticta. Van Tol & Müller (see also Hämäläinen & Müller Distribution (Fig. 108). – Philippines: Luzon (Nueva 1997: 276-277). Thirdly, a third specimen available Viscaya).

181 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

86

84 85

Figures 84-86. Protosticta lepteca sp. n., male [JvT 19212, Luzon, Aurora province, Dinalungan, Mt Anaguao, Alebit river area, 600-900 m, 9-14.iii.1997]. – 84, pronotum, oblique view. – 85, anal appendages, dorsal view. – 86, idem, left lateral view. Scale bar 1 mm.

Protosticta lepteca van Tol, sp. n. mandibles bluish white in basal third, rest brownish (Figs 84-86, 108) black; remaining part of head brownish black with Protosticta sp. n. 25. – Hämäläinen & Müller 1997: 258, 277 metallic shine, finely punctulate; transverse occipital (1 male Quirino province). carina poorly developed, without lateral extremities. Antenna with scapus brown, pedicellus dirty yellow, Type material. – Holotype male [JvT 19218] in RMNH. flagellum castaneous. – ‘Philippines, Luzon Id / Aurora province, Dinalungan Thorax. Pronotum (Fig. 84) predominantly greyish / Mt Anaguao, Alebit river / area, 600-900 m, March 9.- 14. 1997 / R.A. Müller leg u. coll.’ – Paratypes 7 males 1 yellow, with brown markings as follows: medio- female (all Philippine Islands, in RMNH): Luzon. Quirino posterior part of anterior lobe, anterio-median part of province, Maddela, Sulong river, 500-650 m, 26-27.iv.1991 middle lobe, and central parts of tubercles of median (Th. Borromeo) 1 male; Aurora province, Dinalungan, lobe, central part of posterior lobe; lateral lobes fully Mt Anaguao, Alebit river, 600-900 m, 9-14.iii.1997 (R.A. brown; anterior lobe with a paired erect process, flat, Müller) 1 male 1 female; same, Bungo river, 200-500 m, 15- top rounded, approximately as long as median length 18.iii.1997, 5 males. of anterior lobe; posterior lobe with a paired process Diagnosis. – Delicate species, synthorax with as well, fairly close together, narrow and bent at 60˚ narrow longitudinal stripes, especially anterior to anteriad. humeral suture, and over metepisternum posterior Synthorax brownish black with creamish markings to metastigma; differs from other such species with as follows: narrow line over mesepisternum against a missing Ab vein by the following combination of humeral suture, widening both anteriorly and characters (male): long and slender inferior appendage, posteriorly, ending just before posterior margin of pronotum with a pair of short projections on anterior synthorax; mesepimeron with squarish marking lobe, and a paired, triangular, erect process directed against posterior margin of synthorax, dorso- anteriad on posterior lobe; other species have short and anteriorly connected with a fine line against humeral compact appendages and / or more rounded processes suture, starting c. two-fifths from posterior corner on the posterior lobe of the pronotum; the projections of mesokatepisternum; metepisternum with oblong on the anterior corner is also present in Sulcosticta marking against first suture posterior ro metastigma, a striata sp. n., which has distinctly different appendages. longer and broader line along metapleural suture from metastigma to hind margin of metepisternum, the Male [JvT 19128, holotype]. – Head. Labium posteriormost part widening and bent towards middle bluish white, with narrow line along anterior border; of metepisternum; metepimeron with crescent-shaped

182 Chapter 5 Platystictidae of the Philippines

marking against posterior margin. Wings hyaline, Etymology. – Leptekes (Greek): fine-pointed, delicate; venation brown; Px 13 in fore wing, Px 12 in hind for the structure of the projections of the posterior wing; R4+5 arising just distal to subnodus, IR3 half a lobe of the pronotum. An adjective (latinized form of cell distal to that level; Arculus just distal to Ax2; Ab Greek). vein missing; pterostigma reddish brown, width 1.3 Distribution (Fig. 108). – Philippine Islands: Luzon times the height, anterior side convex; cells between (Quirino and Aurora provinces). Costa and R1 posterior to pterostigma undivided. Abdomen with segment 1 brown, segment 2 brown Protosticta plicata van Tol, sp. n. with a paired creamish stripe along anterior margin; (Figs 87-89, 108) segments 3-6 castaneous, anterior one-tenth of each segment creamish white; segment 7 with anterior one- Type material. – Holotype male [JvT 25060] in RMNH: [Philippine Islands]: ‘Kawasan Falls / Badian, Cebu / third bluish white, rest of segment 7 and segments 2/13/2001 / T. Borromeo’. – Paratype (in RMNH): same data 8-10 brownish black; segments 8-10 much wider than as holotype, 1 female. preceding segments, shining. Appendages (Figs 85-86) with superiors creamish white, base darker, slender, Diagnosis. – Relatively large and dark species of the distal two-thirds ventrally projecting on innerside, Philippine Protosticta; males differs from other species inconspicuous tooth; inferiors also slender, basal three- by the structure of the posterior lobe of the pronotum, fifths castaneous, rest pale brown, apical third nearly a distinctly folded projection with the fold globularly straight with very tip bent 90˚ inward. swollen; appendages slender as in P. lepteca sp. n., Measurements. Hind wing 18 mm, abdomen including but posterior lobe of pronotum diagnostic. Female appendages 31 mm. with wing venation as Drepanosticta Laidlaw (Ab vein present), but otherwise very similar to male, including Female [JvT 19213, from Dinalungan, Alebit river]. structure of posterior projections of posterior lobe of – Structure of pronotum and coloration of synthorax pronotum. differ from male of same site as follows: anterior lobe with anterior margin dilated but without projections, Male [JvT 25060, holotype]. – Head. Labium, hind margin of posterior lobe with short and flat mandibles and anteclypeus bluish white, a very posterior tubercles, lateral corners tapered; synthorax fine black line along anterior border of labium and distinctly paler than in male, the creamish white mandibles, hardly visible from above; rest of head markings as follows: narrow line over mesepisternum black, punctulate, with coppery metallic shine; against humeral suture, widening both anteriorly transverse occipital carina poorly developed, no and posteriorly, ending just before posterior margin lateral extremities; antenna with scapus and pedicellus of synthorax; mesepimeron fully creamish white in creamish white, flagellum brown. posterior two-fifths, except for a narrow black line Thorax. Pronotum (Fig. 87) with anterior, median against humeral suture, the pale marking extending and posterior lobes fully creamish white, lateral lobe anteriorly in dorsal half, not reaching anteriormost brownish black; structure of anterior lobe simple, corner, first suture with pale stripe over its full length; posterior lobe with a paired posterior process, halfway metepisternum with oblong marking posterior to bent back nearly 180˚ towards pronotum, the new top metastigma, covering ca. half the space distal to bulbous and very shiny. Synthorax brownish black, metastigma, and a pale stripe in ventral part of with creamish white markings as follows: dorsal carina metepisternum from metastigma to posterior marking; over its full length, a subulate marking in posterior metepimeron with crescent-shaped marking against fourth over humeral suture, a poorly indicated pale posterior margin and pale spot just in front of that. mark on metepisternum. Legs very pale creamish Measurements. Hind wing 18 mm; abdomen 27 mm. white. Wings hyaline, venation brown; Px 13 in

183 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

8789 88

Figures 87-89. Protosticta plicata spec. nov., male [JvT 25060], holotype, Cebu, Badian, Kawasan Falls, 13.ii.2001]. – 87, pronotum, oblique view. – 88, anal appendages, dorsal view. – 89, idem, left lateral view. Scale bar 1 mm.

fore wing 13, Px 12 in hind wing; R4+5 arising at Etymology. – Plicatus (Latin), folded, for the structure subnodus, IR3 half a cell distal to that level; Arculus of the paired process of the posterior lobe of the distal to Ax2; Ab vein absent; pterostigma width 1.2 pronotum. An adjective. times the height, the proximal side somewhat convex, Conservation status. – One of the most threatened but distinctly acutely angulate, distal side less so; cells species of damselflies in the Philippines. The type between Costa and R1 distal to pterostigma undivided. locality is one of three very small remaining patches Abdomen with segment 1 brownish black, segment 2 of forest in Cebu. It has not been recorded from the dark brown, segments 3-6 dark brown with anteriorly other two sites by the research group of Dr. Franz on each segment a creamish white mark, covering Seidenschwarz. Kawasan Falls is not a conservation approximately one-eighth, and three times one-sixth of area, but it be may receive some protection since it is segment-length; creamish white marking on segment a favourite nature travel destination (M. Hämäläinen, 8 more extensive, c. two-fifths segment length, rest personal communication, June 2004). brown, the brown part anteriorly with subulate Distribution (Fig. 108). – Philippine Islands: southern projection; segments 8-10 brownish black, much Cebu (Badian). Only known from type locality. wider than preceding segments. Anal appendages (Figs 88-89) with superiors greyish brown, stoutly built, Sulcosticta van Tol gen. n. somewhat projected on innerside; inferiors surpassing length of superiors, stout at base, distal two-fifths very Diagnosis. – Inferior appendage short and stout, with slender and straight, only the very tip hooked axiad the tip flattened and with shining brown sclerotization, (not dorsad). or more or less bifid, and inner part with or without Measurements. Hind wing 19 mm, abdomen, shining brown sclerotization; synthorax with variegated including appendages, 31 mm. pattern of longitudinal stripes. Wing venation as in Drepanosticta Laidlaw or as in Protosticta Selys. Female. – Similar to the male, but, remarkably, Ab Remarks. – As already explained in the introductory vein present as in Drepanosticta Laidlaw; structure chapter, it is presently, without extensive phylogenetic of posterior lobe of pronotum more or less as male, analysis, impossible to judge which other species will although the folded projection somewhat smaller. ultimately be placed in this genus. The similarity in Mesurements. Hind wing 21 mm, abdomen 31 mm. the structure of the inferior appendage is such that a homoplasy is considered unlikely. Since both Protosticta

184 Chapter 5 Platystictidae of the Philippines

and Drepanosticta are based on characters in wing and posterior lobe dirty yellow, lateral lobe brown; venation, a proper placement of these species in one anterior border of anterior lobe with indistinct triangular of the present genera appeared impossible. It might extremity; median and lateral lobes simple; posterior appear that the variegate coloration of the synthorax lobe with lateral corners erect, triangular, with a sharp, will prove to be a phylogenetically informative constricted top. Synthorax dirty yellow, except for a character for a monophyletic clade, so that also all narrow stripe against dorsal carina, the humeral line, Philippine Protosticta can be placed here. an oblong marking over mesepimeron from just above Type species. – Sulcosticta pallida sp. n. metastigma, and posterior corner of metepimeron. Included species: S. pallida sp. n., S. striata sp. n. and Legs yellow. Wings hyaline, venation brown; Px 14 S. viticula sp. n., and possibly two undescribed species in fore wing, Px 13 in hind wing; origin of R4+5 (damaged male, or only female known). distinctly distal to subnodus; origin of IR3 in middle of Distribution: Luzon. cell distal to subnodus; quadrangle parallel-sided, not widening distally; anal veins complete separated at base; pterostigma rhomboid, width c. 1.5 times the height; Sulcosticta pallida van Tol, sp. n. cells between Costa and R1 complete, undivided. (Figs 90-92, 109) Abdomen. – Segments 1-6 pale brown, with distinct Drepanosticta sp. n. 8. – Hämäläinen & Müller 1997: 257, basal yellow annulae; segments 7-10 dark brown or 277 (Nueva Vizcaya, Nueva Ecija and Quirino provinces). brownish black. Appendages (Figs 91-92) grey, short

Type material. – Holotype male [JvT 22675] in RMNH: and stout, superiors in dorsal view tapering, inner view ‘Philippines, Luzon Id / Nueva Viscaya, Sta Fe / Atbo River, shows large triangular top; inferiors very short, c. two- 550-800 m / June 10, 1991 / Roland A. Müller legit / Coll. thirds the length of superiors, stout, the tip bifurcate. R.A. Müller’. – Paratypes (all Philippines, in RMNH, 7 Measurements. Hind wing 21; abdomen including females): Luzon Id. Data as holotype, 2 females; Nueva appendages (approximate measurements, the Viscaya, Sta Fe, Dalton Pass area, 900 m, 8-17.viii.1991 (A. abdominal segments 8-10 are broken) 31.5. Gorostiza) 1 female; Nueva Ecija, Caraglan, Batching river, 700-850 m, 11.vi.1991 (R.A. Müller) 1 female; Quirino / Female. – As the male, but posterior margin of Aurora boundary, Maddela, Mt Anaguao, 800-1100 m, 9-14. posterior lobe straight without projections, lateral viii.1996 (C.M. Nazareno) 2 female; Aurora, Dinalungan, corners angulate; coloration of synthorax as the male. Mt Anaguao, Bungo river, Lumot creek, 1200-1400 m, Measurements. Hind wing 22-23 mm, abdomen 29-32 20.vi.1997 (M. Dusayen) 1 female. mm. Diagnosis. – The characteristic Y vein ofDrepanostocta not closed, but separate at base, and male appendages Etymology. – Pallidus (Latin), pale; for the coloration very aberrant with inferiors short, the top bifid; the of the synthorax. An adjective. shape of the inferior appendage of the male also present Distribution (Fig. 109). – Philippines: Luzon (Nueva in S. striata sp. n., which differs considerably in the Vizcaya, Nueva Ecija, Quirino and Aurora provinces). shape of the hind lobe of the pronotum. Sulcosticta striata van Tol, sp. n. Male [JvT 22675, holotype]. – Head. Labrum, (Figs 93-95, 109) mandibles and anteclypeus pale, some parts bluish, other parts brownish; a distinct oval depression central Protosticta sp. n. 26. – Hämäläinen & Müller 1997: 258, 277 in labrum; rest of head bronze-black; postoccipital (Luzon). carina with indistinct extremities; parocular suture Type material. – Holotype male [JvT 19219] in RMNH: distinct; antenna incomplete, scapus and pedicellus ‘Philippines, Luzon Id / Quirino/Aurora Boundary / Maddela, pale, the flagellum broken. Mt Anaguao / 9.-14. August 1996, 800-1100 m / Celso M. Thorax. – Pronotum (Fig. 90) with anterior, median Nazareno legit / Coll. Roland A. Müller’. – Paratypes (all

185 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

92 91 90

Figures 90-92. Sulcosticta pallida sp. n., male [JvT 22675, holotype, Luzon, Nueva Viscaya, Sta Fe, Atbo river, 550-800 m, 10.vi.1991]. – 90, pronotum, oblique view. – 91, anal appendages, dorsal view. – 92, idem, left lateral view. Scale bar 1 mm.

Philippine Islands, in RMNH, 7 males 1 female): Luzon. yellow, with brownish black markings as follows: centre Quirino, Maddela, Sierre Madre, Mt Yadanan, Sitio Yadanan, of middle lobe anterior to paired pronotal tubercles, 400-600 m, 15-30.viii.1996 (C.M. Nazareno) 1 male; Aurora lateral lobes in anterior and posterior quarters, province, Dinalungan, Mt Anaguao, Alebit river, 600-900 m, median part of posterior lobe leaving a narrow pale 9-14.iii.1997 (R.A. Müller) 1 male; Aurora Prov., Villa Maria, La-ab river, 360-500 m, 20-24.iii.1997 (R.A. Müller) 5 males line anterior and posterior to this marking. Anterior 1 female. lobe with a paired long tapering process from anterior Other material examined (Philippines, in RMNH). – border of lobe, bending posteriad, the apical half Excluded from type series: Luzon, Camarines Norte, Sn curved nearly 90˚ from body axis; posterior lobe Lorenzo Ruiz, Sitio Bay-Bay, Patag, 400-700 m, 24-25. simple, posterior border only with sides indistinctly ix.1997 (C.M. Nazareno) 1 female [JvT 25064]. produced. Synthorax with complex markings of Diagnosis. – Wing venation as in Protosticta Selys, male brownish black and creamish white: mesepisternum appendages distinct, with inferiors short, the top bifid; brownish black with narrow pale line along humeral the shape of the inferior appendage of the male is also suture, anteriorly above mesokatepisternum slightly present in S. pallida sp. n., which differs considerably widened, line not reaching posterior border of in the shape of the hind lobe of the pronotum, while synthorax; mesokatepisternum brown with pale oblong that species also lacks the projections of the anterior marking near mesepisternum and against mesocoxa; lobe of the pronotum. mesepimeron pale, a dark stripe over humeral suture, a triangular dark marking against first suture anterior Male [JvT 19211, holotype]. – Head. Labium and to level of metastigma, and an irregular oblong line mandibles bluish white, labium with a very narrow posterior to that level; metepisternum pale, the lower brown line along anterior border, and anterior one- half anterior to metastigma brown, a longer line over fourth with brownish shade; anteclypeus bluish white, middle of metepisternum posterior to metastigma, rest of head black, distinctly punctulate with bronze covering approximately one-third of metepisternum, a metallic shine; transverse occipital carina clearly small spot against first suture just anterior to posterior defined, but without lateral extremities; antenna with margin; metakatepisternum brown, metepimeron scapus and pedicellus dirty yellow, flagellum brown. pale, a dark stripe over metapleural suture somewhat Thorax. Pronotum (Fig. 93) predominantly dirty widening in middle, a small dark spot near anterior

186 Chapter 5 Platystictidae of the Philippines

93 95 94

Figures 93-95. Sulcosticta striata sp. n., male [JvT 19220, Luzon, Aurora province, Dinalungan, Mt Anaguao, Alebit river area, 600-900 m, 9-14.iii.1997]. – 93, pronotum, oblique view. – 94, anal appendages, dorsal view. – 95, idem, left lateral view. Scale bar 1 mm.

corner, a dark oblong marking in lower posterior inferiors wide and flattened, somewhat hollow, shorter corner. Legs very pale. Wings hyaline, venation than superiors, the tip emarginate with inner-tooth brown; Px 13 in fore wing, Px 12 (or 12½) in hind longer and curved dorsad. wing; R4+5 arising distinctly posterior to subnodus, Measurements. Hind wing 19 mm, abdomen including especially in fore wing, IR3 arising at c. one-third of appendages c. 30 mm. length of cell posterior to that level; Arculus distinctly posterior to Ax2; quadrangle not widening posteriorly, Female. – The only female assigned to this species is not even in fore wing; anal bridge absent (as in teneral and in rather poor condition; the processes Protosticta); pterostigma approximately as wide as of the anterior lobe of pronotum present, but much high, anterior and posterior sides convex in fore wing, shorter than in male. proximal side more straight, but angulate, in hind Measurements. Hind wing 19 mm, abdomen including wing; cells distal to pterostigma undivided. appendages c. 30 mm. Abdomen generally pale coloured with darker parts of segments 2-6 very pale brown; segment 1 dark brown, Note. – The female from Quirino province mentioned lateral parts and an annule against segment 2 creamish in Hämäläinen & Müller (1997: 277) under note white; segment 2 creamish with large, crescent-shaped 26 [JvT 26990] is not conspecific and mentioned brown marking over full segment, enclosing medio- separately below under Sulcosticta sp. B. The female posteriorly a dark spot; segments 3-6 pale brown, all mentioned above and excluded from the type series segments both anteriorly and posteriorly paler with [JvT 25064] is teneral and in poor condition. The annules of segments dark; segment 7 with anterior projections of the posterior lobe of the pronotum are two-fifths bluish white, rest of segment 7 and segments somewhat larger than in the other female assigned to 8-10 brownish black; segments 8-10 distinctly wider this species. This specimen is preliminarily identified as than preceding segments. Appendages (Figs 94-95) pale S. striata sp. n. and short; base of superiors dorsally somewhat darker, Etymology. – Striatus (Latin), striped, for the base relatively slender and both appendages widely coloration of the synthorax. An adjective. separate, the distal half turned 90˚ and thus dorso- Distribution (Fig. 109). – Philippine Islands: Luzon ventrally oriented, the lower and distal margins sharp; (Quirino, Aurora).

187 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

98 97 96

Figures 96-98. Sulcosticta viticula spec. nov., male [JvT 25062, holotype. Luzon, Zambales province, Masinloc, Mt Coto, Tal-tal, 400-700 m, 1-8.iv.1997]. – 96, pronotum, oblique view. – 97, anal appendages, dorsal view. – 98, idem, left lateral view. Scale bar 1 mm (for figures 97-98).

Sulcosticta viticula van Tol, sp. n. Thorax. Pronotum (Fig. 96) predominantly dirty (Figs 96-98, 109) yellow, with dark brown markings as follows: centre

Type material. – Holotype male [JvT 25062]: Philippines, of middle lobe anterior to paired pronotal tubercles, Luzon Id., Zambales Prov., Masinloc, Mt Coto, Tal-tal, 400- the lateral lobes, and median part of posterior lobe 700 m, 1-8.iv.1997 (C.M. Nazareno) [in RMNH]. – Paratype leaving a narrow pale line posterior to this marking. (in RMNH): same data as holotype, 1 female. Anterior lobe erect with a paired projection near lateral corners, the projections short and erect; posterior Diagnosis. – Small and slender species with distinct lobe simple, posterior border laterally with rounded pale longitudinal stripes over synthorax; Ab vein projections, approximately as long as median line of absent; male differs from other Philippine species with posterior lobe. Synthorax with complex markings of this combination of characters (Protosticta annulata, P. brownish black and creamish white: mesepisternum lepteca and Sulcosticta striata) by structure of inferior brownish black with narrow pale line along humeral appendage: a subterminal process approximately suture, anteriorly above mesokatepisternum slightly diameter of appendage directed axiad, then posteriad, widened, line not reaching posterior border of the tip brownish black and directed abaxiad; top of synthorax; mesokatepisternum brown with small appendix rounded; combination of tips of appendage pale marking above mesocoxa; mesepimeron pale, a and large subterminal process superfically looking dark stripe over humeral suture, an oblong triangular as bifid top. This combination otherwise only in the brownish black marking against first suture, anteriorly unnamed species from Polillo (Sulcosticta sp. A). from halfway between mesocoxa and metastigma, tapering posteriorly to four-fifths of length of Male [JvT 25062, holotype]. – Head. Labium bluish mesepimeron; metepisternum pale, the lower three- white, anterior border with brownish shade only; quarters anterior to metastigma brown, a longer line mandibles with anterior two-thirds pale brown, rest over middle of metepisternum posterior to metastigma, bluish white; anteclypeus bluish white; rest of head covering approximately one-third of metepisternum, black, distinctly coriaceous with bronze metallic shine; narrow brown stripe against metapleural suture; transverse occipital carina distinct, but without lateral metakatepisternum brown, metepimeron brown, a extremities; antenna with scapus and pedicellus dirty small pale spot in posterior corner. Legs very pale. yellow, flagellum brown. Wing hyaline, venation brown; Px 11 in fore wing, Px

188 Chapter 5 Platystictidae of the Philippines

11 in hind wing; R4+5 arising at subnodus, IR3 arising Unplaced specimens halfway length of cell posterior to that level; Arculus Drepanosticta indet. A distinctly posterior to Ax2; quadrangle somewhat widening posteriorly in fore wing; anal bridge absent A female with coloration of synthorax as D. moorei, but (as in Protosticta); pterostigma 1.4 times as wide as without conspicuous projections on pronotum. Hind high, anterior side acutely angulate; posterior side wing 24 mm, abdomen ca. 33 mm. Transverse occipital more rectangular; cells between Costa and R1 distal to carina without lateral extremities. pterostigma very few and undivided. Abdomen rather pale coloured, with darker parts of Material examined (Philippines, in RMNH). – Luzon Id., segments 2-6 very pale brown; segment 1 dark brown; Aurora Prov., Villa Maria, La-ab River, 360-500 m, 20-24. segment 2 brown with crescent-shaped pale markings iii.1997 (R.A. Müller) 1 female [JvT 26989]. in anterior lower corners; segments 3-6 pale brown, all segments with anterior one-tenth to one-seventh Drepanosticta indet. B dirty white; segment 7 with anterior two-fifths white Material examined (Philippines, in RMNH). – Panaon with a brown annule anteriorly, rest of segment 7 and Id., San Francisco, Batong Lapd, viii.1988 (W. Catal) 1 segments 8-10 brownish black, shiny; segments 8-10 female [JvT 18618], and second label ‘Drepanosticta lestoides distinctly wider than preceding segments. Appendages (Brauer), M. Hämäläinen det’. (Figs 97-98) pale and short; base of superiors dorsally slightly darker; base relatively slender and the pair A female with distinct lateral extremities of transverse widely separate, curved inward, distal half turned 90˚, occipital carina, pair of long projections on anterior club-shaped, with inconspicuous dorsal tooth, ventral lobe of pronotum and smoothly rounded posterior side straight; inferiors wide and flattened, top rounded, lobe of pronotum, synthorax with white stripe over distal two-fifths with complicated curled structure: a metepisternum presumably represents a new species. subterminal, stoutly built process directed axiad, then Hind wing 20 mm, abdomen ca. 30 mm. posteriad, the tip brownish black and turned abaxiad. Measurements. Hind wing 17 mm, abdomen including Sulcosticta sp. A appendages c. 26 mm. Material examined. – “Polillo, Philipp., 20.VIII.1915. G. Boettcher”, 1 male (in SMFD) [JvT 26975]. Female. – Generally as the male, but projections of anterior lobe of pronotum missing, lateral projections Diagnosis. – Small and slender species with distinct of posterior lobe of pronotum much smaller; pale pale longitudinal stripes over synthorax; Ab vein markings on mesepimeron and metepisternum dististly absent; male differs from other Philippine species larger in male, but generally of similar shape. with this combination of characters by structure of Measurements. Hind wing 18 mm; abdomen 29 mm. inferior appendages: short and tapering, distal one- fifth compressed and shining brown, the tip recurved Etymology. – Viticula (Latin), tendril; for the shape of dorso-axiad, tip ending in a comb of teeth. Apart from the inward directed process of the inferior appendage. appendages very similar to S. viticula sp. n. A noun in apposition. Distribution (Fig. 109). – Philippine Islands: Luzon Male [JvT 26975]. – Head. Labium bluish white, (Zambales). anterior border with brownish shade only; mandibles with anterior two-thirds pale brown, rest bluish white; anteclypeus bluish white; rest of head black, distinctly coriaceous with bronze metallix shine; transverse

189 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

occipital carina distinct, but without lateral extremities; annule; segments 3-6 pale brown, all segments with antenna broken. anterior one-tenth to one-seventh paler (indistinct, but Thorax. Pronotum predominantly dirty yellow, with this possibly due to poor preservation); segment 7 with brown markings as follows: centre of middle lobe anterior two-fifths pale brown, rest of segment 7 and anterior to paired pronotal tubercles dark brown; segments 8-10 brownish black, shiny; segments 8-10 lateral lobes castaneous; median part of posterior distinctly wider than preceding segments. Appendages lobe brown, leaving a narrrow pale line anterior and (damaged and right superior appendage missing) pale posterior to marking. Anterior lobe erect with a paired and short, base relatively slender and the pair originally projection near lateral corners, the projections short widely separate, curved inward, distal half turned 90˚ and erect; posterior lobe simple, posterior border dorso-ventrad, club-shaped with inconspicuous dorsal laterally with rounded projections, approximately tooth, ventral side curved, the tip distinctly angulate; as long as median line of posterior lobe. Synthorax inferiors wide, short, stout, and tapering, distal one- with complex markings of brownish black and fifth compressed and shining brown, the tip recurved creamish white: mesepisternum brownish black with dorso-axiad, tip ending in a comb of short teeth. narrow pale line along humeral suture, anteriorly Measurements. Hind wing 17 mm; abdomen with anal above mesokatepisternum broadly widened, the line appendages ca. 26 mm. itself just reaching posterior border of synthorax; mesokatepisternum brown with small pale marking Note. – Due to the poor state of conservation I have above mesocoxa; mesepimeron pale, a narrow dark refrained from naming this specimen. Further studies stripe of humeral suture; metepisternum pale, the lower on Polillo island may reveal new material of this anterior three-quarters pale brown, a narrow brown phylogenetically interesting taxon. stripe against metapleural suture; metakatepisternum Distribution. – Polillo Island (off eastern Luzon). castaneous; metepimeron castaneous, a small pale spot in anterior corner, posterior part somewhat paler than Sulcosticta sp. B rest. Legs very pale yellowish white. Wings hyaline, venation brown; Px 12 in fore wing, Px 11 in hind Material examined. – Philippines, Luzon Id. Quirino, Maddela, Sierre Madre, Mt Yadanan, Sitio Yadanan, 400-600 wing; R4+5 arising well distal to subnodus, IR3 arising m, 15-30.viii.1996 (C.M. Nazareno), 1 female [JvT 26990]. halfway length of cell posterior to that level; Arculus arising at Ax2; quadrangle hardly widening posteriorly; One female from Mt Yadanan in RMNH possibly anal bridge absent (as in Protosticta); pterostigma 1.3 belongs to the genus Sulcosticta as defined here, times as wide as high, subrectangular, anterior and with pale thoracic markings and an Ab and Ac vein posterior veins convex; cells between Costa and R1 present, but separate at wing margin. Coloration of distal to pterostigma very few and undivided. the synthorax is clearly different from S. pallida sp. n., Abdomen (broken and partly repaired). Rather pale while the presence of the Ab vein, plus the structure coloured, with darker parts of segments 2-6 very pale of the pronotum is distinct from S. striata. The latter brown; segment 1 castaneous with posterior pale species was also recorded from Mt. Yadanan, so a yellow annule; segment 2 pale with crescent-shaped further study of this interesting complex of species is dark markings from lower latero-posterior corners needed when more material becomes available. Hind to middle of segment anteriorly, a brown posterior wing 24 mm, abdomen ca. 33 mm.

Figures 99-102. Distribution of Drepanosticta species in the Philippines. – 99, Drepanosticta aries group. – 100, Drepanosticta belyshevi group. – 101, Drepanosticta moorei group. – 102, Drepanosticta lymetta group.

190 Chapter 5 Platystictidae of the Philippines

99 100

D. acuta D. belyshevi D. aries D. flavomaculata D. krios D. trachelocele D. rhamphis

101 102

D. luzonica D. clados D. moorei D. furcata D. hermes D. lymetta D. taurus

191 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

Acknowledgements References

The names of Roland A. Müller and Dr. Matti Asahina, S., 1984. A list of the Odonata from Thailand. Part Hämäläinen will remain associated with the III. Platystictidae. – Kontyû 52: 585-595. advancement of our knowledge of the odonates of Brauer, F., 1868. Dritter Bericht über die von Herrn G. Semper mitgetheilten, von dessen Bruder auf den the Philippines during the last 25 years of the 20th Philippinen gesammelten Neuropteren und Beschreibung century. Roland A. Müller (St Gallen, Switzerland) einer neuen Libellen-Gattung. – Verhandlungen K. K. brought together a huge and extremely rich collection zoologisch-botanischen Gesellschaft Wien 18: 541-558. between c. 1980 and 1997. Dr Matti Hämäläinen Cowley, J., 1936. Descriptions of three new species of (Espoo, Finland) sorted, studied and identified all Drepanosticta (Odonata) from the Philippine Islands. – Transactions of the Royal Entomological Society of London material. I am grateful to both colleagues for entrusting 85: 157-168, figs 1-22. their collections to Museum Naturalis, and leaving the Gassmann, D. & M. Hämäläinen, 2002. A revision of the scientific study of some parts to me. Matti Hämäläinen Philippine subgenus Risiocnemis (Igneocnemis) Hämäläinen had expertly sorted the specimens, and I have (Odonata: Platycnemididae). – Tijdschrift voor followed most, but not all, of his opinions on species Entomologie 145: 213-266. delimitation; of course all decisions shall be considered De Jong, R. & C.G. Treadaway, 1993. The Hesperiidae (Lepidoptera) of the Philippines. – Zoologische mine. Dr. Hämäläinen showed me for study the Verhandelingen 288: 1-125, figs 1-139. specimens of SMFD he had on loan, and also kindly Dickerson, R.E., 1928. Distribution of life in the Philippines. reviewed a draft of this paper. – Macmillan, New York: 309 pp. Thanks are due to the staff artists of the National Hämäläinen, M., 1991. Drepanosticta belyshevi spec. nov. Museum of Natural History, Leiden, the Netherlands, from the Philippines (Zygoptera: Platystictidae). – Odonatologica 20: 65-68. Bas (S.B.) Blankevoort, Erik Jan Bosch and Inge Hämäläinen, M., 1997. Stenagrion petermilleri spec. nov. from M. van Noortwijk for preparing such attractive Palawan, the Philippines (Zygoptera: Coenagrionidae). – illustrations of this remarkable group of damselflies. Odonatologica 26: 473-475. The specimens ofProtosticta plicata sp. n. were Hämäläinen, M., 1999. Drepanosticta jurzitzai spec. nov., collected by Dr. F. Seidenschwarz and Mr. Borromeo a new damselfly from southeastern Thailand (Zygoptera: during a biodiversity survey on Cebu, and were Platystictidae). – Odonatologica 28: 421-423. Hämäläinen, M., 2000. Risiocnemis seidenschwarzi spec. nov., kindly put at my disposal for this paper via Dr. M. an endangered damsefly from Tabunan forest in Cebu, the Hämäläinen. Mr. Reagan J. Villanueva (Davao City, Philippines (Odonata: Platycnemididae). – Entomologische Philippines) presented the specimens of Drepanosticta Berichten 60: 46-49. clados sp. n. to the Leiden Museum, and guided me to Hämäläinen, M. & R.A. Müller, 1997. Synopsis of the some interesting sites in northern Mindanao in April Philippine Odonata, with lists of species from forty islands. – Odonatologica 26: 249-315, figs 1-7. 2004. Laidlaw, F.F., 1917. A list of the dragonflies recorded from the Finally, I am grateful to Steve Brooks (BMNH), James Indian Empire with special reference to the collection of K. Liebherr (CUIC) and Philip D. Perkins (MCZC) the Indian Museum. Part II. The family Agrioninae. A. The for loan of material or information on material in the sections Podolestes, Platycnemis, Platysticta and Protoneura. collections under their care. – Records of the Indian Museum 13: 321-348, figs 1-17, plates xiii-xv. Lieftinck, M.A., 1937. Descriptions and records of South-East Asiatic Odonata, – Treubia 16: 55-119. Lieftinck, M.A., 1939. Sur quatre espèces nouvelles de Platystictidae de l’Inde orientale (Odonates). – Revue française d’Entomologie 6: 144-154, figs 1-4.

Figures 103-106. Distribution of Drepanosticta species in the Philippines. – 103, Drepanosticta megametta group. – 104, Drepa- nosticta lesttoides Brauer. – 105, Drepanosticta mylitta Cowley. – 102, Drepanosticta pistor sp. n.

192 Chapter 5 Platystictidae of the Philippines

103 104

D. lestoides D. centrosaurus D. megametta

105 106

D. mylitta D. pistor

193 van Tol. – Phylogeny and biogeography of the Platystictidae (Odonata)

Lieftinck, M.A., 1961. Philippine Zoological Expedition Mémoires couronnés l’Académie royale de Belgique 38: 1946-1947. New and interesting Odonata from the sep. 1-233. Philippines. – Fieldiana (Zoology) 42 (10): 119-149, figs Selys Longchamps, E. de, 1891. Additions aux odonates des 1-52. Philippines. – Anales de la Sociedad Española de Historia Lieftinck, M.A., 1965. Some Odonata of the genus Natural 20: 209-218. Drepanosticta Laidlaw, chiefly from the Malay Peninsula Semper, G., 1892. Die Schmetterlinge der philippinischen (Platystictidae). – Zoologische Mededelingen 40 (21): 171- Inseln. 7. Lieferung: 271-380. Wiesbaden. 186, figs 1-15. van Tol, J., 2000. The Odonata of Sulwesi and adjacent Lieftinck, M.A., 1974. Dragonflies collected by the Noona islands. Part 5. The genusProtosticta Selys (Platystictidae). – Dan Expedition in the southwestern Philippine Islands Tijdschrift voor Entomologie 143: 221-266, figs 1-113. (Insecta, Odonata). – Steenstrupia 3: 111-147, figs 1-26. van Tol, J. & R.A. Müller, 2003. Forest damselflies of the Needham, J. G. & M.K. Gyger, 1939. The Odonata of the Philippines, their evolution and present status, with the Philippines, II. – Philippine Journal of Science 70: 239- description of Drepanosticta moorei spec. nov. from Luzon 314 + plates 11-22. (Zygoptera: Platystictidae). – Odonatologica 32: 39-45, Needham, J.G. & M.K. Gyger, 1941. More Odonata from figs 1-5. the Philippines. – Philippine Journal of Science 74: 141- Vane-Wright, R.I., 1990. The Philippines – Key to the 151 + plate 1. biogeography of Wallacea? p. 19-34. – In: W.J. Knight & Orr, A.G., 2003. A guide to the dragonflies of Borneo. Their J.D. Holloway (Eds), Insects and the rain forests of South East identification and biology. – Natural History Publications, Asia (Wallacea). Entomological Society of London: i-iv + Kota Kinabalu: i-x + 1-195, figs 1-210, plates 1-25. 1-343. Selys Longchamps, E. de, 1882. Odonates de Philippines. – Watson, J.A.L. & A.F. O’Farrell, 1991. Odonata: p. 294-310. Anales de la Sociedad Española de Historia Natural 11: sep In: I.D. Naumann et al., The insects of Australia. A textbook 1-32 + 1 plate. for students and research workers. Volume 1. Second ed. Selys Longchamps, E. de, 1885. Programme d’une Melbourne University Press: 542 pp. révision des Agrionines. – Compte rendu de la Société Wilson, K.D.P., 1997. The Platystictidae of Hong Kong and entomologique de Belgique 29: cxlvi-cxlvii. Guangdong, with descriptions of a new genus and two new Selys Longchamps, E. de, 1886. Révision du synopsis des species. – Odonatologica 26: 53-63, figs 1-31. Agrionines. Première partie, comprenant les légions Wilson, K.D.P. & G.T. Reels, 2001. Odonata of Hainan, Pseudostigma, Podagrion, Platycnemis et Protoneura. – China. – Odonatologica 30: 145-208, figs 1-173.

194 Chapter 5 Platystictidae of the Philippines

107 108

D. aurita D. ceratophora P. annulata D. malleus P. lepteca D. myzouris P. plicata D. paruatia D. quadricornu

109

S. pallida S. striata S.viticula

Figures 107-109. Distribution of Platystictidae in the Philippines. – 107, Distribution of D. aurita sp. n., D. ceratophora Lieftinck, D. malleus sp. n., D. myzouris sp. n., D. paruatia sp. n., D. quadricornu sp. n. – 108, Distribution of the species of Protosticta Selys. – 109. Distribution of the species of Sulcosticta gen. n. 195